Evolution of the ovenbird-woodcreeper assemblage

(Aves: Furnariidae) �/ major shifts in nest architecture and

adaptive radiation

Martin Irestedt, Jon Fjeldsa and Per G. P. Ericson

Irestedt, M., Fjeldsa, J. and Ericson, P. G. P. 2006. Evolution of the ovenbird-woodcreeper assemblage (Aves: Furnariidae) �/ major shifts in nest architecture andadaptive radiation. �/ J. Avian Biol. 37: 260�/272

The Neotropical ovenbirds (Furnariidae) form an extraordinary morphologically andecologically diverse passerine radiation, which includes many examples of species thatare superficially similar to other passerine birds as a resulting from their adaptations tosimilar lifestyles. The ovenbirds further exhibits a truly remarkable variation in nesttypes, arguably approaching that found in the entire passerine clade. Herein we presenta genus-level phylogeny of ovenbirds based on both mitochondrial and nuclear DNAincluding a more complete taxon sampling than in previous molecular studies of thegroup. The phylogenetic results are in good agreement with earlier molecular studies ofovenbirds, and supports the suggestion that Geositta and Sclerurus form the sisterclade to both core-ovenbirds and woodcreepers. Within the core-ovenbirds severalrelationships that are incongruent with traditional classifications are suggested. Amongother things, the philydorine ovenbirds are found to be non-monophyletic. Themapping of principal nesting strategies onto the molecular phylogeny suggests cavitynesting to be plesiomorphic within the ovenbird�/woodcreeper radiation. It is alsosuggested that the shift from cavity nesting to building vegetative nests is likely to havehappened at least three times during the evolution of the group. We suggest that theshifts in nest architecture within the furnariine and synallaxine ovenbirds have served asan ecological release that has facilitated diversification into new habitats and newmorphological specializations.

M. Irestedt (correspondence) and P. G. P. Ericson, Department of VertebrateZoology and Molecular Systematics Laboratory, Swedish Museum of Natural History,P.O. Box 50007, SE-104 05 Stockholm, Sweden. E-mail: martin.irestedt@nrm.se.J. Fjeldsa, Vertebrate Department, Zoological Museum, University of Copenhagen,Universitetsparken 15, DK�/2100 Copenhagen Ø, Denmark.

The Neotropical ovenbirds (Furnariidae) exhibit a

unique morphological and ecological variation among

passerine birds (Leisler 1977, Vaurie 1980, Remsen

2003). The group includes many examples of species

that exhibit remarkable superficial morphological

similarities with other passerine birds that only are

distantly related to ovenbirds, for instance the cinclodes

species (Cinclodes ) resembling muscicapine thrushes

(Turdidae) or dippers (Cinclidae), miners (Geositta )

resembling larks (Alaudidae), earthcreepers (Upucerthia )

resembling thrashers (Toxostoma , Mimidae) and various

synallaxine genera resembling various Old World

grass warblers (‘‘Sylviidae’’) or creepers (Certhiidae).

Also in their placement and structure of the nest the

ovenbirds exhibit an extraordinary diversity. In fact,

the variation in nest types in this family has been

suggested to approach that found in the entire passerine

clade (Sick 1993, Collias 1997, Zyskowski and Prum

1999, Remsen 2003). Nevertheless, breeding in ‘‘closed’’

nest chambers, whether placed inside a cavity or in

the vegetation, seems to unite all ovenbirds (and

woodcreepers).

# JOURNAL OF AVIAN BIOLOGY

JOURNAL OF AVIAN BIOLOGY 37: 260�/272, 2006

260 JOURNAL OF AVIAN BIOLOGY 37:3 (2006)

Clades of passerine birds differ greatly in their degree

of morphological divergence (Ricklefs 2003). Remark-

able radiations on small and species-poor landmasses,

such as islands, are often interpreted as a consequence of

competition for a limited range of food types (e.g., Grant

1986). The variation in the ovenbird clade may represent

another model, given their vast distribution in South and

Central America and the great diversity of other kinds of

birds. ‘‘Key innovations’’ may represent another possible

explanation for non-random variation in diversity

among clades. The relative success of passerine birds

has for example been postulated to be promoted by the

complexity of the passerine foot (Raikow and Bledsoe

2000) or by the ability to build nests from a wide variety

of materials (Olson 2001). The ability of ovenbirds to

construct nests of a tremendous architectural diversity

could thus be a possible explanation for their successful

colonization of a wide range of habitats, along with a

special modification of the kinetic properties of the bill,

which allowed them to extract food that was otherwise

hidden in complex vegetation structures (Fjeldsa et al.

2005).

Monophyly of the ovenbirds and woodcreepers is

inferred from their shared, unique syrinx morphology

(Ames 1971), and has also been supported by molecular

data (Sibley and Ahlquist 1990, Irestedt et al. 2002,

Chesser et al. 2004, Fjeldsa et al. 2005). Mostly based on

overall similarity ovenbirds have traditionally been

divided into three major groups, Furnariinae, Synallax-

inae and Philydorinae (i.e. Hellmayr 1925, Vaurie 1971,

1980; see Sibley and Ahlquist 1990 for a historical

review), with the woodcreepers as their closest relatives.

However, Feduccia (1973) noted cranial similarities

between certain woodcreepers and the philydorine oven-

birds, and suggested that the woodcreepers had evolved

from a philydorine ovenbird. Several recent DNA

analyses strongly support Feduccia?s hypothesis that

woodcreepers are nested within the ovenbirds, but reject

a close affinity to the philydorines. Instead, it is

suggested that the leaftossers (Sclerurus ) and miners

(Geositta ) are basal to both woodcreepers and core

ovenbirds, and that these latter two groups are recipro-

cally monophyletic (Irestedt et al. 2002, Chesser 2004,

Fjeldsa et al. 2005). The molecular studies also suggest

other relationships at odds with traditional classifica-

tions of ovenbirds: the placement of Pseudoseisura

among the synallaxines and of Lochmias among the

furnariines (as opposed to their traditional placement

among philydorine ovenbirds), and Xenops in a basal

position in the woodcreeper radiation (instead of among

philydorines) (Fjeldsa et al. 2005). These studies ob-

viously raise doubts about the monophyly of the

‘‘philydorine’’ ovenbirds.

Herein, we present a molecular phylogenetic hypoth-

esis of ovenbird relationships based on an expanded

taxonomic sample and data from two nuclear introns,

myoglobin intron 2 and G3PDH intron 11, and the

mitochondrial cytochrome b gene. Based on these

phylogenetic results we will reassess the evolution of

nest-building strategies of ovenbirds.

Materials and methods

Taxon sampling, amplification and sequencing

The ingroup in this study includes 48 ovenbirds (repre-

senting 40 out of 55 genera recognized by Remsen 2003)

and 9 woodcreepers. The ovenbirds were chosen to

represent all major subgroups previously suggested

(Vauri 1971, 1980, Feduccia 1973, Raikow 1994, Ridgely

and Tudor 1994, Zimmer and Isler 2003). In addition,

we included several species that have proven difficult to

place in either of these subgroups. A selection of

woodcreepers was also included, as recent molecular

data have shown this group to be nested within oven-

birds (Irestedt et al. 2002, Chesser 2004, Fjeldsa et al.

2005). As outgroups we used two representatives of the

family Rhinocryptidae (Pteroptochos tarnii and Scyta-

lopus spillmanni ) and one of the family Formicariidae

(Chamaeza meruloides ), which form the sister clade to

the Furnariidae (Irestedt et al. 2002, Chesser et al. 2004).

Sample identifications and GenBank accession numbers

are given in Table 1.

The complete myoglobin intron 2, the complete

glyceraldehydes-3-phosphodehydrogenase (G3PDH) in-

tron 11, and 999 bp from the cytochrome b gene have

been sequenced (see Fjeldsa et al. 2003, and Irestedt et

al. 2002 for sequencing procedures). For each gene and

taxon, multiple sequence fragments were obtained by

sequencing with different primers. These sequences were

assembled to complete sequences with SeqMan IITM

(DNASTAR Inc.). Positions where the nucleotide could

not be determined with certainty were coded with the

appropriate IUPAC code. Due to the low number of

insertions in the introns the combined sequences could

easily be aligned by eye. All gaps have been treated as

missing data in the analyses. No insertions, deletions,

stop or nonsense codons were observed in any of the

cytochrome b sequences.

Certhiaxis cinnamomea produced a partly unreadable

sequence for the G3PDH intron 11, and the PCR

product from this taxon was cloned. The cloning,

amplification and sequencing were done with the

TOPO TA Cloning† Kit (Invitrogen Life Technologies),

using the manufacturer’s primers and protocol. In one of

the two clones studied, an autapomorphic insertion was

found at the positions where the sequences from the

uncloned PCR-products became unreadable. Thus, this

insertion likely explains the reading problems in the

latter. In the phylogenetic analysis of the G3PDH intron

11 both clones were included, but as these two clones

grouped together, the G3PDH partition in Certhiaxis

JOURNAL OF AVIAN BIOLOGY 37:3 (2006) 261

Table 1. Samples used in the study. Family and subfamily names follow the classification of Remsen (2003). Abbreviations: AHMN�/American Museum of Natural History, New York;ANSP�/Academy of Natural Sciences of Philadelphia; NRM�/Swedish Museum of Natural History; ZMUC�/Zoological Museum of the University of Copenhagen. References: (1)Irestedt et al. (2001), (2) Irestedt et al. (2004b), (3) Ericson et al. (2002), (4) Fjeldsa et al. (2005), and Olson et al. (2005).

Species Subfamily/family Sample No. Cytochrome b Myoglobin G3PDH

Geositta rufipennis Furnariidae: Furnariinae ZMUC S290 AY590042 (ref. 4) AY590052 (ref. 4) AY590062 (ref. 4)Geositta tenuirostris Furnariidae: Furnariinae ZMUC S292 AY590043 (ref. 4) AY590053 (ref. 4) AY590063 (ref. 4)Upucerthia jelskii Furnariidae: Furnariinae ZMUC S439 AY065700 (ref. 1) AY065756 (ref. 1) AY590064 (ref. 4)Cinclodes fuscus Furnariidae: Furnariinae ZMUC S220 AY590044 (ref. 4) AY590054 (ref. 4) AY590065 (ref. 4)Furnarius leucopus Furnariidae: Furnariinae ZMUC 125590 AY996351 (ref. 5) AY996345 (ref. 5) AY996357 (ref. 5)Furnarius cristatus Furnariidae: Furnariinae NRM 966772 AY064279 (ref. 3) AY064255 (ref. 3) AY590066 (ref. 4)Limnornis curvirostris Furnariidae: Synallaxinae USNM B2735 AY996353 (ref. 5) AY996346 (ref. 5) AY996359 (ref. 5)Limnoctites rectirostris Furnariidae: Synallaxinae USNM B14895 AY996352 (ref. 5) AY996347 (ref. 5) AY996358 (ref. 5)Phleocryptes melanops Furnariidae: Synallaxinae USNM B2734 AY996354 (ref. 5) AY996348 (ref. 5) AY996360 (ref. 5)Aphrastura spinicauda Furnariidae: Synallaxinae ZMUC 134531 AY998188 AY998225 AY998206Leptasthenura pileata Furnariidae: Synallaxinae ZMUC S338 AY590045 (ref. 4) AY590055 (ref. 4) AY590067 (ref. 4)Schizoeaca harterti Furnariidae: Synallaxinae ZMUC S2398 AY998189 AY998226 AY998207Oreophylax moreirae Furnariidae: Synallaxinae ZMUC S128816 AY998190 AY998227 AY998208Schoeniophylax phryganophilus Furnariidae: Synallaxinae NRM 947184 AY998191 AY998228 AY998209Synallaxis ruficapilla Furnariidae: Synallaxinae NRM 956643 AY065707 (ref. 1) AY065763 (ref. 1) AY590068 (ref. 4)Synallaxis scutata Furnariidae: Synallaxinae ZMUC 125635 AY998192 AY998229 AY998210Hellmayrea gularis Furnariidae: Synallaxinae ZMUC 124846 AY998193 AY998230 AY998211Cranioleuca sulphurifera Furnariidae: Synallaxinae USNM B17199 AY996350 (ref. 5) AY996344 (ref. 5) AY996356 (ref. 5)Cranioleuca pyrrhophia Furnariidae: Synallaxinae NRM 966821 AY065708 (ref. 1) AY065764 (ref. 1) AY590069 (ref. 4)Cranioleuca albicapilla Furnariidae: Synallaxinae ZMUC 124797 AY996349 (ref. 5) AY996343 (ref. 5) AY996355 (ref. 5)Certhiaxis cinnamomeus Furnariidae: Synallaxinae NRM 937358 AY998194 AY998231 AY998212, AY998213Asthenes cactorum Furnariidae: Synallaxinae ZMUC S150 AY065705 (ref. 1) AY065761 (ref. 1) AY590070 (ref. 4)Asthenes urubambensis Furnariidae: Synallaxinae ZMUC S172 AY998195 AY998232 AY998214Phacellodomus ruber Furnariidae: Synallaxinae NRM 947206 AY590046 (ref. 4) AY590056 (ref. 4) AY590071 (ref. 4)Anumbius annumbi Furnariidae: Synallaxinae NRM 966903 AY065709 (ref. 1) AY065765 (ref. 1) AY590072 (ref. 4)Coryphistera alaudina Furnariidae: Synallaxinae NRM 966910 AY065710 (ref. 1) AY065766 (ref. 1) AY590073 (ref. 4)Metopothrix aurantiaca Furnariidae: Synallaxinae NRM 569302 AY998224Xenerpestes singularis Furnariidae: Synallaxinae ANSP 4370 AY998205Premnornis guttuligera Furnariidae: Philydorinae ZMUC 128014 AY998196 AY998233 AY998215Premnoplex brunnescens Furnariidae: Philydorinae ZMUC 124927 AY998197 AY998234 AY998216Margarornis squamiger Furnariidae: Philydorinae ZMUC S1112 AY065703 (ref. 1) AY065759 (ref. 1) AY590074 (ref. 4)Pseudoseisura lophotes Furnariidae: Philydorinae NRM 976799 AY998199 AY998236 AY998218Pseudocolaptes boissonneautii Furnariidae: Philydorinae ZMUC 124935 AY998198 AY998235 AY998217Berlepschia rikeri Furnariidae: Philydorinae ZMUC S1214 AY590047 (ref. 4) AY590057 (ref. 4) AY590075 (ref. 4)Anabacerthia striaticollis Furnariidae: Philydorinae ZMUC 124673 AY998200 AY998237 AY998219Syndactyla rufosuperciliata Furnariidae: Philydorinae ZMUC124972 AY998201 AY998238 AY998220Philydor atricapillus Furnariidae: Philydorinae NRM 937334 AY065702 (ref. 1) AY065758 (ref. 1) AY590076 (ref. 4)Thripadectes flammulatus Furnariidae: Philydorinae ZMUC S428 AY065701 (ref. 1) AY065757 (ref. 1) AY590077 (ref. 4)Automolus leucophthalmus Furnariidae: Philydorinae NRM 937251 AY590048 (ref. 4) AY590058 (ref. 4) AY590078 (ref. 4)Hylocryptus erythrocephalus Furnariidae: Philydorinae ZMUC 124862 AY998202 AY998239 AY998221Sclerurus mexicanus Furnariidae: Philydorinae ZMUC S1443 AY590049 (ref. 4) AY590059 (ref. 4) AY590079 (ref. 4)Sclerurus scansor Furnariidae: Philydorinae NRM 937258 AY065715 (ref. 1) AY065772 (ref. 1) AY590080 (ref. 4)Lochmias nematura Furnariidae: Philydorinae ZMUC S2577 AY065699 (ref. 1) AY065755 (ref. 1) AY590081 (ref. 4)Heliobletus contaminatus Furnariidae: Philydorinae ZMUC 127191 AY998203 AY998240 AY998222Xenops minutus Furnariidae: Philydorinae ZMUC S451 AY590050 (ref. 4) AY590060 (ref. 4) AY590082 (ref. 4)Xenops rutilans Furnariidae: Philydorinae ZMUC S452 AY590051 (ref. 4) AY590061 (ref. 4) AY590083 (ref. 4)Megaxenops parnaguae Furnariidae: Philydorinae ZMUC 125605 AY998204 AY998241 AY998223Pygarrhichas albogularis Furnariidae: Philydorinae AMNH PRS1128 AY065704 (ref. 1) AY065760 (ref. 1) AY590084 (ref. 4)

26

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cinnamomea in the combined analysis is represented by

the consensus sequence from both clones.

Phylogenetic inference and model selection

Bayesian inference and Markov chain Monte Carlo

(MCMC) were used for estimating phylogenetic hypoth-

esis from our DNA data (see recent reviews by Huel-

senbeck et al. 2001, Holder and Lewis 2003). The models

for nucleotide substitutions used in the analyses were

selected for each gene individually by using Akaike

Information Criterion (AIC, Akaike, 1973) and the

program MrModeltest (Nylander 2002) in conjunction

with PAUP* (Swofford 1998).

The posterior probabilities of trees and parameters in

the substitution models were approximated with MCMC

and Metropolis coupling using the program MrBayes

(Ronquist and Huelsenbeck 2003). Analyses were per-

formed for both the individual gene partitions and the

combined data set. In the analysis of the combined data

set the models selected for the individual gene partition

were used, but the topology was constrained to be the

same. One cold and three incrementally heated chains

were run for 2.5 million generations, with a random

starting tree. Trees were sampled every 100th genera-

tions, and the trees sampled during the burn-in phase

(i.e., before the chain had reached its apparent target

distribution) were discarded. Two runs, starting from

different, randomly chosen trees, were made to ensure

that the individual runs had converged on the same

target distribution (Huelsenbeck et al. 2002). After

checking for convergence, final inference was made

from the concatenated output from the two runs.

Results

Variation in the molecular data set

The sequences obtained ranged from 667 bp (Helioble-

tus ) to 716bp (Chamaeza ) in the myoglobin intron 2,

and from 287 bp (Chamaeza ) to 410 bp (Margarornis ) in

the G3PDH intron 11. Most indels observed in the two

introns were autapomorphic and mainly found in

particularly variable and repetitive regions. However,

some synapomorphic indels were observed when map-

ping the data onto the tree topologies obtained from the

Bayesian analyses of the combined data set. In the

myoglobin intron 2, all ovenbirds and woodcreepers lack

28 bp present in the outgroup (Scytalopus also lack 12 of

these bp but shares the remaining 16 bp with the other

outgroup taxon); the woodcreeper representatives share

a deletion of 12 bp; Pteroptochos tarnii and Scytalopus

spillmanni share a deletion of 13 bp and an insertion of

10 bp; Sclerurus mexicanus and Sclerurus scansor share

a deletion of 4 bp; and Philydor atricapillus andTab

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JOURNAL OF AVIAN BIOLOGY 37:3 (2006) 263

Heliobletus contaminatus share a deletion of 14 bp. In

the relatively more indel-rich G3PDH intron 11 indels

with different lengths sometimes overlap. However, if

these are treated as independent events the following

synapomorphic indels are found; Geositta rufipennis and

tenuirostris share a deletion of 23 bp; Deconychura ,

Dendrocincla and Sittasomus share two deletions of 34

and 1 bp, respectively; Furnarius cristatus and leucopus

share a deletion of 19 bp; Xenops minutus and rutilans

share a deletion of 1 bp; Pteroptochos tarnii and

Scytalopus spillmanni share a deletion of 3 bp;

all ovenbirds and woodcreepers share an extra basepair

lacking in the outgroup and in the representatives

of the genera Geositta and Sclerurus ; Anabacerthia ,

Automolus, Heliobletus, Hylocryptus, Megaxenops,

Philydor and Syndactyla share an insertion of 1 bp

(Thripadectes has an autapomorphic deletion in

that region); and Heliobletus and Philydor share an

insertion of 9 bp.

A few indels were also found to be incongruent

with the phylogenetic tree obtained from the analysis

of the combined data set. These were generally found

in the most variable regions and some of the single

basepair insertions actually consist of different bases.

However, one of these indels in myoglobin consists of

a 4 bp deletion shared between Certhiaxis cinnamomea

and Schoeniophylax phryganophila . The remaining,

phylogenetically incongruent indels were found to in

the G3PDH intron 11 and includes a 1 bp insertion in

Cinclodes, Deconychura and Premnoplex ; a 1 bp inser-

tion in Asthenes cactorum and Furnarius cristatus ; a 1 bp

deletion in Automolus and Thripadectes ; a 1 bp deletion

in Pteroptochos and Glyphorynchus ; a 10 bp deletion

in Drymornis and Xiphorhynchus ; and a 3 bp insertion in

Glyphorynchus, Campylorhamphus, Drymornis, Nasica ,

Xiphocolaptes and Xiphorhynchus.

The observed, pairwise distances (p-distances) range

between 0.3% (Limnoctites rectirostris and Cranioleuca

sulphurifera , Synallaxis scutata and Synallaxis rufica-

pilla , Cranioleuca pyrrhophia and Cranioleuca albica-

pilla ) and 8.8% (Scytalopus spillmanni and Synallaxis

scutata ) in myoglobin, 0.0% (Xenops minutus and

rutilans ) and 13.2% (Berlepschia rikeri and Scytalopus

spillmanni , Berlepschia rikeri and Chamaeza meruloides )

in G3PDH, and between 2.4% (Lochmias nematura and

Upucerthia jelskii ) and 20.3% (Chamaeza meruloides and

Synallaxis ruficapilla , Pteroptochos tarnii and Nasica

longirostris ) in cytochrome b.

Model selection and phylogenetic relationships

The priori selection of substitution models supported

that the GTR�/I�/G model had the best fit for the

cytochrome b , and the myoglobin partitions, while the

GTR�/G model was selected for the G3PDH intron 11.

These models were used in the Bayesian analyses of

the individual genes as well as in the combined analysis.

After discarding the burn-in phase the inference for

the cytochrome b gene was based on a total of 48,400

samples from the posterior, while the inference

for myoglobin and G3PDH introns were based on

47,800 samples and 47,500 samples for the combined

data set. For the phylogenetic inference, the mode of

the posterior distribution of topologies was presented

as a majority-rule consensus tree from each analysis

(Figs. 1 and 2).

The trees obtained from the Bayesian analyses of

the individual gene partitions (Fig. 1) shows a similar

degree of resolution (although the tree obtained from

the cytochrome b partition is slightly more resolved).

The gene trees are also overall topologically similar

and the non-congruent relationships are mostly at short

nodes with rather modest posterior probabilities. Phylo-

genetic relationships congruently supported by all

three genes includes a strong support for a sister

relationship between the genera Geositta and Sclerurus ;

a clade consisting of the Cranioleuca representatives

and Limnoctites rectirostris ; a core foliage-gleaners

clade (Anabacerthia , Syndactyla , Megaxenops, Helioble-

tus, Philydor, Automolus, Hylocryptus, and Thripa-

dectes ); and a clade consisting of traditional

Furnariinae ovenbirds (Furnarius, Cinclodes and Upu-

certhia ) plus the genera Lochmias, Phleocryptes and

Limnornis.

A few conflicting topologies between the individual

gene trees are supported by rather solid posterior

probabilities, but there is no indication that one gene

tree should be more different than the others. Examples

of strong topological conflicts include the relative

position of Limnoctites rectirostris. While the G3PDH

tree gives a strong support placing Limnoctites recitros-

tris with Cranioleuca albicapilla , both the myoglobin

and cytochrome b trees placed it with C. sulphurifera .

Another example is in the relative positions of Certhiaxis

cinnamomea and Synallaxis scutata , where the topology

of the G3PDH tree differs from those in the myoglobin

and cytochrome b trees. There are also examples where

the cytochrome b tree or the myoglobin tree supports

relationships different from what suggested by the

G3PDH tree. For example, the cytochrome b tree

supports a basal position of Xenops minutus and Xenops

rutilans relative to all other ovenbirds except the genera

Geositta and Sclerurus, while both the G3PDH and

myoglobin trees suggest that Xenops instead is basal

within the woodcreeper clade. The analysis of the

combined data set results in a tree (Fig. 2) that is better

resolved than the individual gene trees, although topo-

logically overall similar to them. Overall the combined

tree tends to support relationships that are supported by

a majority of the gene partitions.

264 JOURNAL OF AVIAN BIOLOGY 37:3 (2006)

Discussion

Phylogenetic resolution

The overall topological similarity between the individual

gene trees (Fig. 1) support that the tree obtained in the

analysis of the combined data set (Fig. 2) is an overall

good estimate of the generic relationships among oven-

birds. Nevertheless, the topological disagreements that

do exist between the gene trees might cause some

concern about the accuracy of certain parts of the

combined tree.

It has been postulated that trees based on the

mitochondrial genome has a better chance to correctly

estimate avian phylogenies than single nuclear genes

(Moore 1995, Moore and DeFilippis 1997), but exam-

ples exist where mitochondrial trees yield phylogenetic

estimates that are at odd with both nuclear data and the

relationships that could be expected based on biogeo-

graphy or morphological data (Degnan 1993, Alstrom

and Odeen 2002). Furthermore, even though the ob-

served topological conflicts could be genuine, i.e., that

the different genomic partitions used herein have differ-

ent phylogenies (reviewed in e.g., Moore 1995, Maddison

1997, Mindell 1997), the conflicts might also be due to

inaccuracies of the substitution models, or in the

methods used to infer the trees. Without further evidence

from, for example, DNA sequence data obtained from

other linkage groups, morphological or behavioral

characters, it seems virtually impossible to discriminate

between individual gene trees. As the combined tree is

based on all gene partitions it tends to exhibit relation-

ships supported by a majority of these. Consequently, we

believe that the combined tree overall is a better

Fig. 1. The 50% majority ruleconsensus trees obtained from theBayesian analyses of the individualgenes. A) the tree obtained from theanalyses of G3PDH (glyceraldehydes-3-phosphodehydrogenase) intron 11data set, B) the tree obtained from theanalyses of the myoglobin intron 2 dataset, and C) tree obtained from theBayesian analyses of the cytochrome bdata set. Posterior probability valuesare indicated to the right of the nodes.

Cranioleuca albicapillaLimnoctites rectirostris

Cranioleuca pyrrhophia

Cranioleuca sulphurifera

Metopothrix aurantiaca

Certhiaxis cinnamomeus cl 2Certhiaxis cinnamomeus cl 1

Schoeniophylax phryganophilusSynallaxis ruficapilla

Asthenes cactorumPseudoseisura lophotes

Synallaxis scutataOreophylax moreirae

Schizoeaca hartertiAsthenes urubambensis

Aphrastura spinicaudaLeptasthenura pileata

Anumbius annumbiCoryphistera alaudina

Hellmayrea gularisPhacellodomus ruber

Premnoplex brunnescensLimnornis curvirostris

Phleocryptes melanopsLochmias nematura

Cinclodes fuscusUpucerthia jelskii

Furnarius cristatusFurnarius leucopus

Premnornis guttuligeraPseudocolaptes boissonneautiiMargarornis squamiger

Pygarrhichas albogularisAnabacerthia striaticollisSyndactyla rufosuperciliata

Megaxenops parnaguaeHeliobletus contaminatus

Philydor atricapillusAutomolus leucophthalmus

Hylocryptus erythrocephalusThripadectes flammulatus

Berlepschia rikeri

Nasica longirostrisXiphocolaptes major

Campylorhamphus trochilirostrisDrymornis bridgesii

Xiphorhynchus erythropygius

Glyphorynchus spirurus

Dendrocincla tyranninaSittasomus griseicapillus

Deconychura longicauda

Xenops minutusXenops rutilans

Sclerurus mexicanusSclerurus scansor

Geositta rufipennisGeositta tenuirostris

Pteroptochos tarniiScytalopus spillmanni

Chamaeza meruloides

1.000.99

1.00

0.72

0.991.00

1.00

0.520.63

0.501.00

0.90

0.69

0.95

0.960.79

0.78

0.70

0.77

0.961.00

0.96

1.000.98

0.97

1.00

1.00

0.98

1.000.99

1.00

0.98

1.00

0.78

0.790.72

0.74

0.59

0.66

1.00

0.86

0.78

0.99

1.00

1.00

1.00

A)

JOURNAL OF AVIAN BIOLOGY 37:3 (2006) 265

phylogenetic estimate than any of the individual gene

trees. In fact, we found relationships based on the

individual genes to be in conflict with those of the

combined tree mainly where nodes are short and poster-

ior probabilities modest (B/0.95).

The combined phylogeny is in many respects incon-

gruent with the traditional division of ovenbirds into

three major groups, Furnariinae, Synallaxinae and

Philydorinae (i.e. Hellmayr 1925, Vaurie 1971, 1980).

Many recent molecular studies of other passerine bird

groups (i.e. Madagascan songbirds, see Cibois et al.

2001; antbirds, see Irestedt et al. 2004a) have shown that

traditional classifications, based on overall similarities,

often overlooked natural relationships, and the present

result is therefore no great surprise. The overall good

phylogenetic agreement between the results presented

herein and other studies of ovenbirds based on DNA-

sequences partly obtained from other genes (Irestedt

et al. 2002, Chesser 2004) are, on the other hand, a

strong indication that our combined phylogeny is a good

estimate of the generic relationships of ovenbirds.

The finding that Geositta and Sclerurus are basal

positioned to both the woodcreepers and remaining

ovenbirds are, for example, also conclusively supported

by the b-fibrinogen intron 7 (Chesser 2004). In addition,

recently revealed morphological characters also support

several of these novel molecular hypotheses. Fjeldsa

et al. (2005) demonstrate that Geositta and Sclerurus

share similar skull morphologies with the so called

‘‘transitory woodcreepers’’ (Feduccia 1973), and differ

from the skull morphologies found in other ovenbirds.

An even more unexpected example, exposed by Fjeldsa

et al. (2005), is that Xenops shares with Glyphorhynchus

a unique functional system for hammering in wood. Also

a different interpretation of Clench’s (1995) study of the

pterylosis patterns of woodcreepers and ovenbirds sup-

port this relationship, as Xenops and woodcreepers

shares a lower number of feathers in the pars dorsalis

Fig. 1 (Continued )Cranioleuca sulphuriferaLimnoctites rectirostris

Cranioleuca albicapillaCranioleuca pyrrhophia

Synallaxis ruficapillaSynallaxis scutata

Schoeniophylax phryganophilusCerthiaxis cinnamomeus

Pseudoseisura lophotesOreophylax moreirae

Schizoeaca hartertiAsthenes urubambensis

Anumbius annumbiPremnornis guttuligera

Hellmayrea gularis

Limnornis curvirostrisPhleocryptes melanops

Cinclodes fuscusUpucerthia jelskii

Furnarius cristatusFurnarius leucopus

Lochmias nematura

Aphrastura spinicaudaCoryphistera alaudina

Asthenes cactorumPhacellodomus ruber

Leptasthenura pileata

Pseudocolaptes boissonneautii

Automolus leucophthalmus

Thripadectes flammulatusHylocryptus erythrocephalus

Anabacerthia striaticollisSyndactyla rufosuperciliataMegaxenops parnaguae

Heliobletus contaminatusPhilydor atricapillus

Margarornis squamigerPremnoplex brunnescens

Pygarrhichas albogularis

Berlepschia rikeri

Drymornis bridgesiiXiphorhynchus erythropygius

Campylorhamphus trochilirostris

Xenops minutusXenops rutilans

Deconychura longicaudaSittasomus griseicapillus

Dendrocincla tyranninaNasica longirostris

Xiphocolaptes majorGlyphorynchus spirurus

Geositta rufipennisGeositta tenuirostris

Sclerurus mexicanusSclerurus scansor

Pteroptochos tarniiScytalopus spillmanni

Chamaeza meruloides

1.00

1.00

0.990.90

1.00

1.00

1.00

0.73

1.000.83

0.99

1.00

1.00

1.000.93

1.00

1.00

0.90

1.00

0.950.89

0.780.96

0.860.68

0.76

1.00

0.99

1.00

1.00

0.56

0.781.00

0.590.73

0.941.00

1.001.00B)

266 JOURNAL OF AVIAN BIOLOGY 37:3 (2006)

than found in ovenbirds. Nevertheless, other morpholo-

gical characters are sometimes in conflict with the results

presented herein. Thus, the ‘‘Margarornis assemblage’’,

as defined from characters from the hindlimb muscu-

lature (Rudge and Raikow 1992), is suggested to be non-

monophyletic by our molecular data. However, Irestedt

et al. (2004b) and McCracken et al. (1999) have found

similar conflicts between molecular data and morpho-

logical characters from the hindlimb, suggesting that

hindlimb characters may often be under adaptive selec-

tion and less useful in phylogenetic reconstructions. This

might also be case for several other morphological

characters (that have been used to infer relationships

among ovenbirds and woodcreepers) that are in conflict

with the present molecular phylogeny.

As a consequence of our much denser taxon sampling

than in previous DNA-based phylogenies of ovenbirds,

several taxa with contested affinity can now be placed,

some traditional groupings can for the fist time be

corroborated by molecular data, while other groups

could be rejected or modified. While the Furnariinae

and Synallaxinae in large parts remains intact, although

with somewhat different generic compositions, the

philydorine ovenbirds are unlikely to be monophyletic

(see also Fjeldsa et al. 2005): four ‘‘philydorine’’ clades

could be recognized; (1) a core foliage-gleaners clade

(Anabacerthia , Syndactyla , Megaxenops, Heliobletus,

Philydor, Automolus, Hylocryptus, and Thripadectes ),

(2) a clade with Margarornis, Premnoplex and Pygar-

rhichas, (3) a clade with Premnornis and Pseudocolaptes,

and (4) a clade for the aberrant Berlepschia . Except for

the Premnornis and Pseudocolaptes clade, these foliage-

gleaner clades are positioned basal to traditional furnar-

iines and synallaxines, but the combined phylogeny

suggests that they are separated by fairly long inter-

nodes. Most noticeable is the position of Premnornis and

Pseudocolaptes, which strongly suggests that the Phily-

dorinae is an unnatural group.

Fig. 1 (Continued )

Cranioleuca albicapillaCranioleuca pyrrhophia

Cranioleuca sulphuriferaLimnoctites rectirostris

Xenerpestes singularisAsthenes cactorum

Schoeniophylax phryganophilusSynallaxis scutata

Synallaxis ruficapilla

Certhiaxis cinnamomeusPseudoseisura lophotes

Oreophylax moreiraeSchizoeaca harterti

Asthenes urubambensisAnumbius annumbi

Coryphistera alaudinaHellmayrea gularis

Phacellodomus ruberLeptasthenura pileata

Premnornis guttuligeraPseudocolaptes boissonneautii

Limnornis curvirostrisPhleocryptes melanops

Lochmias nematuraFurnarius cristatus

Furnarius leucopus

Cinclodes fuscusUpucerthia jelskii

Aphrastura spinicauda

Berlepschia rikeri

Anabacerthia striaticollisSyndactyla rufosuperciliata

Philydor atricapillus

Megaxenops parnaguaeHeliobletus contaminatus

Automolus leucophthalmusThripadectes flammulatus

Hylocryptus erythrocephalus

Premnoplex brunnescensPygarrhichas albogularis

Margarornis squamiger

Xenops minutusXenops rutilans

Campylorhamphus trochilirostrisDrymornis bridgesii

Xiphorhynchus erythropygiusNasica longirostris

Xiphocolaptes majorDeconychura longicauda

Sittasomus griseicapillusDendrocincla tyrannina

Glyphorynchus spirurusSclerurus mexicanus

Sclerurus scansorGeositta rufipennis

Geositta tenuirostrisPteroptochos tarnii

Scytalopus spillmanniChamaeza meruloides

1.00

0.76

1.001.00

1.00

0.84

0.96 1.00

0.64

1.000.80

0.59

0.66

1.00

0.95

1.00

0.94

0.69

1.00

0.53

1.000.74

0.620.90

1.000.56

0.830.64

1.000.801.00

0.61

0.78

0.96

0.93

0.640.99

1.001.00

1.00

1.00

0.64

1.00

0.97

0.62 0.64

1.00

1.00

1.00C)

JOURNAL OF AVIAN BIOLOGY 37:3 (2006) 267

Ecological shifts and morphology

The new phylogenetic groupings might seem surprising

in view of the general similarities in external morphology

between ‘‘philydorine’’ ovenbirds (even though certain

genera such as Margarornis and Pygarrhichas are quite

divergent from the typical philydorines). However, based

on our combined phylogeny (Fig. 2), it is obvious that

deeper branches (except for Geositta ) comprise forest

birds of a fairly uniform appearance.

The most parsimonious solution of the ovenbird

radiation is thus an early forest diversification, leading

to divergence of woodcreepers, a large radiation of

foliage-gleaners, most of which obtained their food by

probing and prying in internodes and among masses of

dead leaves and debris suspended among vines and

branches (Fjeldsa et al. 2005), and lineages that specia-

lized to search food in epiphyte-clad cloud-forest and

shrubbery of the rising Andean mountains. Thus the

terminal radiations of Furnariinae and Synallaxinae

ovenbirds (generally adapted to more xeric life zones

and more opened landscapes) would represent more

drastic shifts of environment.

Although the Furnariinae and Synallaxinae clades

remain largely intact in the combined molecular tree,

some of the suggested relationships are novel. Most

noticeable is the placement of reedhaunters, with Lim-

nornis forming a clade with Phleocryptes, close to

Lochmias, within the Furnariinae ovenbirds, and Lim-

noctites being placed among the Synallaxinae ovenbirds.

The relationships of all these have been obscure, but the

suggested relationship between Phleocryptes and Lim-

nornis is supported by their similar nest architecture

(Zyskowski and Prum 1999). These relationships are

discussed in further detail by Olson et al. (2005).

Other forest taxa that have been difficult to place are

Megaxenops and Heliobletus, which now can be placed,

with high confidence, among the philydorine foliage-

gleaners. Cytochrome b and G3PDH data, respectively,

also provide good evidence for placing Xenerpestes and

Fig. 2. The 50% majority ruleconsensus tree obtained from theanalyses of the combined data set(G3PDH intron 11, the myoglobinintron 2 and the cytochrome b datasets). Posterior probability values areindicated to the right of the nodes.

Cranioleuca sulphuriferaLimnoctites rectirostris

1.00

Cranioleuca albicapillaCranioleuca pyrrhophia1.00

1.00

Synallaxis ruficapillaSynallaxis scutata

1.00

Schoeniophylax phryganophilus0.97

Certhiaxis cinnamomeus1.00

0.98

Asthenes cactorumPseudoseisura lophotes

0.93

1.00

Oreophylax moreiraeSchizoeaca harterti

1.00

Asthenes urubambensis1.00

0.99

Anumbius annumbiCoryphistera alaudina

1.00

Hellmayrea gularis0.70

1.00

Phacellodomus ruber

0.89

Leptasthenura pileata0.98

Aphrastura spinicauda0.96

Limnornis curvirostrisPhleocryptes melanops

1.00

Lochmias nematura0.99

Cinclodes fuscusUpucerthia jelskii

1.000.68

Furnarius cristatusFurnarius leucopus

1.001.00

Premnornis guttuligeraPseudocolaptes boissonneautii

1.00

0.90

1.00

Margarornis squamigerPremnoplex brunnescens

0.69

Pygarrhichas albogularis0.99

0.74

Anabacerthia striaticollisSyndactyla rufosuperciliata

0.98

Megaxenops parnaguae1.00

Heliobletus contaminatusPhilydor atricapillus

1.001.00

Automolus leucophthalmusHylocryptus erythrocephalus

0.97

Thripadectes flammulatus1.00

1.00

Berlepschia rikeri

1.00

Campylorhamphus trochilirostrisDrymornis bridgesii

0.95

Xiphorhynchus erythropygius1.00

Nasica longirostrisXiphocolaptes major

1.00

1.00

Deconychura longicaudaSittasomus griseicapillus

0.95

Dendrocincla tyrannina1.00

1.00

Glyphorynchus spirurus1.00

Xenops minutusXenops rutilans

1.00

1.00

1.00

Sclerurus mexicanusSclerurus scansor

1.00

Geositta rufipennisGeositta tenuirostris

1.00

1.00

1.00

Pteroptochos tarniiScytalopus spillmanni

1.00

Chamaeza meruloides

268 JOURNAL OF AVIAN BIOLOGY 37:3 (2006)

Metopothrix among the synallaxines, near the Cranio-

leuca group, and it is therefore likely that Siptornis and

the odd, new-described Acrobatornis (Pacheco et al.

1996) also belong here.

It is finally interesting to see that the large genus

Asthenes, which is mainly associated with open montane

habitats, may not be a natural taxon. Further studies are

needed to see how many Asthenes species (and Thripo-

phaga softtails) that fall outside the main group

(represented here by A. cactorum ).

Environmental factors clearly put morphological

constraints on birds. Certain morphologies are repeat-

edly observed in obviously unrelated passerine species

that occupy forested habitats, while other morphologies

are shared by species living in open habitats. Not

surprisingly, the large and ecologically diverse

ovenbird�/woodcreeper radiation includes many exam-

ples of morphological parallels to distantly related

passerine birds with similar adaptations (Remsen

2003). Given two independent cases of cavity nesters

colonizing open habitats in the ovenbird�/woodcreeper

radiation (Geositta and the Furnariinae clade) and that

shifts from cavity nesting to building vegetative nests

have independently happened at least three times within

the ovenbird�/woodcreeper radiation (see below), it is

not surprising to also find several examples of parallel

morphological evolution within this radiation. However,

other instances of similar-looking taxa, for example the

‘‘philydorine’’ Premnoplex and Premnornis, may not be

explained by convergent evolution. Instead, the phylo-

geny suggests that the deep nodes in the ovenbird�/

woodcreeper radiation mainly consisted of forest birds

that nested in cavities and that often resemble each other

in size, plumage and shape. The superficial similarities

between various ‘‘philydorine’’ lineages may thus be due

to their shared retention of a plesiomorphic, ‘‘philydor-

ine’’ appearance.

Shifts in nest architecture as innovations in the

diversification of ovenbirds

As mentioned, ovenbirds exhibit an extraordinary di-

versity in nest placement and structure, including the

construction of various types of nests in cavities and

crevices, massive ‘‘houses’’ in clay, and vegetative

structure, which in some cases attain gigantic dimensions

and also serve as dormitories for family groups (Zys-

kowski and Prum 1999, Remsen 2003). Mapping of the

general nesting strategies (as coded by Zyskowski and

Prum 1999) onto the combined phylogeny indicates that

cavity nesting is plesiomorphic within ovenbirds (Fig. 3),

a hypothesis that accords with the frequent use of cavity-

nesting among the closest relatives of Furnariidae

(Ridgely and Tudor 1994, Krabbe and Schulenberg

2003) and with the phylogenetic results by Zyskowski

and Prum (1999). The woodcreepers often use cavities

rather low in the trees, or even subterranean (as

Sclerurinae), and have simple nest-cups, and we suggest

that this was the ancestral condition in the Furnariidae.

The shift from cavity nesting to building vegetative

nests is a rather rare evolutionary event in birds, but our

data suggest that it has happened at least three times in

the Furnariidae: One time early in the evolution of the

synallaxine clade, another time in the ancestor of some

genera within the enlarged furnariine clade, and a

third time within the ‘‘philydorine’’ Pygarrhichas �/

Premnoplex �/Margarornis clade (Fig. 3). However, note

that the synallaxine, furnariine, and Pygarrhichas �/Pre-

mnoplex �/Margarornis clades are rather terminal

lineages that share a common ancestor in our phylogeny.

An alternative explanation might thus be that this

common ancestor sometimes placed a reduced vegetative

nest in cavities and built a more advanced vegetative nest

placed in the open at other times, as do Premnoplex and

Leptasthenura . The pre-disposed ability to vary nest

architecture may then have been the major source for the

nest patterns we find within these clades today. Note also

that our phylogeny lacks Eremobius, a genus close to

Upucerthia based on morphology that builds a stick-nest

placed in the open.

It is noticeable that species that built enclosed

vegetative nests within cavities are basal in all the

lineages where this major shift in nests architecture

took places, suggesting that the shift from cavity nesting

to building enclosed vegetative nests is a gradual process.

Lochmias, which is positioned basally in the phylogeny

to the vegetative nesters Limnornis and Phleocryptes

within the furnariine clade, builds a domed vegetative

nest within a cavity. Within the ‘‘philydorine’’

Pygarrhichas �/Premnoplex �/Margarornis clade, Premno-

plex often places its domed moss nest in a cavity, and

that of Margarornis is often placed under a limb or a

rock. Aphrastura , which constitutes the most basal

Fig. 3. Distribution of nest habits within the ovenbird-woodcreeper radiation. Principal nest strategies (nest in cavity and/or domedvegetative structure) have been mapped on the tree obtained from the analysis of the combined molecular data set. Data on nesthabits are mainly from Zyskowski and Prum (1999), and for a few taxa from Remsen (2003). Black branches indicate cavity nesting,blue branches represent the habit of building enclosed vegetative nests placed in the open, and green branches indicate that enclosedvegetative nests are placed within cavities. Note also that the black branches to the representatives of Furnarius have been markedwith blue stripes as it is difficult to tell whether the complex, domed Furnarius nest, built of clay and dung with admixed hairs andplant fibres, should be regarded as a case of cave-nesting (as coded by Zyskowski and Prum 1999) or as homologous with a domedvegetative nest. The asterisk after the names Premnoplex brunnescens and Leptasthenura pileata indicates that these taxa sometimesplace their vegetative nests in cavities and sometimes in the open. Two asterisks mark taxa for which the nest habits are insufficientlyknown. Note that Xenerpestes singularis and Metopothrix aurantiaca have tentatively been added to the tree based on their relativeposition in the cytochrome b and G3PDH trees, respectively.

JOURNAL OF AVIAN BIOLOGY 37:3 (2006) 269

Cranioleuca sulphuriferaLimnoctites rectirostrisCranioleuca albicapillaCranioleuca pyrrhophiaXenerpestes / MetopothrixSynallaxis ruficapillaSynallaxis scutataSchoeniophylax phryganophilusCerthiaxis cinnamomeusAsthenes cactorumPseudoseisura lophotesOreophylax moreiraeSchizoeaca hartertiAsthenes urubambensisAnumbius annumbiCoryphistera alaudinaHellmayrea gularis**Phacellodomus ruberLeptasthenura pileata*Aphrastura spinicaudaLimnornis curvirostrisPhleocryptes melanopsLochmias nematuraCinclodes fuscusUpucerthia jelskiiFurnarius cristatusFurnarius leucopusPremnornis guttuligera**Pseudocolaptes boissonneautiiMargarornis squamigerPremnoplex brunnescens*Pygarrhichas albogularisAnabacerthia striaticollisSyndactyla rufosuperciliataMegaxenops parnaguae**Heliobletus contaminatus**Philydor atricapillusAutomolus leucophthalmusHylocryptus erythrocephalusThripadectes flammulatusBerlepschia rikeri**Campylorhamphus trochilirostrisDrymornis bridgesiiXiphorhynchus erythropygiusNasica longirostrisXiphocolaptes majorDeconychura longicaudaSittasomus griseicapillusDendrocincla tyranninaGlyphorynchus spirurusXenops minutusXenops rutilansSclerurus mexicanusSclerurus scansorGeositta rufipennisGeosittatenuirostrisPteroptochos tarniiScytalopus spillmanniChamaeza meruloides

Fig. 3 (Continued )

270 JOURNAL OF AVIAN BIOLOGY 37:3 (2006)

branch in the synallaxine clade, builds a cup-shaped nest

that partially covers the wall of the cavity. Leptasthe-

nura , the next branch in the synallaxine clade, builds a

domed vegetative nest that often is placed within a cavity

(Zyskowski and Prum 1999).

Building a closed nest structure inside a cavity would

immediately seem to represent unnecessary extra labour,

but may have served a function of securing less humid

conditions inside the nest. It is noteworthy that the

change in nest construction happened in lineages in-

habiting montane environments, and it is worth noting in

this respect that also Andean tapaculos (Scytalopus )

often build a closed nest of moss inside cavities. It is

unfortunate, for this interpretation, that the nest of

Hellmayrea , which inhabits humid Andean forest, is still

undescribed. Lochmias breeds near streams in montane

forests, thus in a humid environment, and Leptasthenura

spp. inhabit a broad habitat spectrum from humid tree-

lines to semiarid highlands. However, other taxa repre-

senting deeper synallaxine branches, Phacellodomus,

Coryphistera and Anumbius, mainly inhabit drier scrub

and savanna, where they assemble enormous masses of

interwoven thorny twigs, containing a system of tunnels,

and a nest chamber. It is therefore not fully clear under

what environmental conditions the shifts in nest-con-

struction happened.

The fact that the terminal synallaxine and furnariine

radiations comprise no less than 115 species and 38

species, respectively, compared with 119 species in all the

remaining lineages of Furnariidae (incl. Dendrocolapti-

nae), suggests that the shift in nest architecture may have

served as an innovation spurring further diversification.

This includes adaptations to new environments. Natural

cavities or ground patches suitable for excavating tunnels

are often in limited supply in open landscapes (Remsen

2003), and the ability to build own vegetative nest

obviously gives a competitive advantage in such environ-

ments. In case of marsh habitats, home of Phleocryptes,

Limnornis and Limnoctites, there is no possibility to find

a cavity or excavate a tunnel. In the Andean cloud-

forests a small closed nest structure can easily be hidden

in the enormous masses of epiphytes that are often

found, which may have played a role among the

synallaxines, and could also explain why Margarornis

and Premnoplex build domed moss nests.

It is noticeable that several of the nest characteristics

of ovenbirds, as coded by Zyskowski and Prum (1999),

become synapomorphies for subclades of taxa when

mapped onto our combined phylogeny. Examples are the

Sphagnum nests of Schizoeaca and Oreophylax , the

thatch placed over the camber in Certhiaxis, Schoenio-

phylax and Synallaxis, the roof adornments in Anum-

bius and Coryphistera , and the pensile nest-type of

Cranioleuca and possibly Xenerpestes (Remsen 2003, p.

319). Other nest characteristics exhibit a large degree of

convergent evolution. However, a phylogeny based on a

denser taxon sampling, as well as a more information

about nest architecture in critical groups, is needed to get

a full understanding of the evolution of nesting strategies

in ovenbirds.

Acknowledgements �/ Tissue and blood samples were mainlyobtained from the Zoological Museum of Copenhagen (withdata collecting supported by the Danish Research Councils) andthe Swedish Museum of Natural History (collected incollaboration with the Museo Nacional de Historia Naturaldel Paraguay, San Lorenzo). Important samples have also beenobtained from the American Museum of Natural History, NewYork, Academy of Natural Sciences of Philadelphia, and theNational Museum of Natural History, Smithsonian Institution.Mari Kallersjo provided logistic support and advice for thework at the Molecular Systematics Laboratory at the SwedishMuseum of Natural History, and Pia Eldenas and DarioZuccon is thanked for practical support at the laboratory.Bernd Leisler and an anonymous reviewer are thanked forvaluable comments on the manuscript. The Swedish ResearchCouncil (grant no. 621�/2004�/2913 to P.E.) funded thelaboratory work.

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(Received 3 January 2005, revised 8 April 2005, accepted9 April 2005.)

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