MORPHOLOGICAL AND ECOLOGICAL EVOLUTION IN OLD AND NEW

WORLD FLYCATCHERS

A dissertation presented to

the faculty of

the College of Arts and Sciences of Ohio University

In partial fulfillment

of the requirements for the degree

Doctor of Philosophy

Clay E. Corbin

August 2002

This dissertation entitled

MORPHOLOGICAL AND ECOLOGICAL EVOLUTION IN OLD AND NEW

WORLD FLYCATCHERS

BY

CLAY E. CORBIN

has been approved for

the Department of Biological Sciences

and the College of Arts and Sciences by

Donald B. Miles

Associate Professor, Department of Biological Sciences

Leslie A. Flemming

Dean, College of Arts and Sciences

CORBIN, C. E. Ph.D. August 2002. Biological Sciences.

Morphological and Ecological Evolution in Old and New World Flycatchers (215pp.)

Director of Dissertation: Donald B. Miles

In both the Old and New Worlds, independent clades of sit-and-wait insectivorous

birds have evolved. These independent radiations provide an excellent opportunity to test

for convergent relationships between morphology and ecology at different ecological and

phylogenetic levels. First, I test whether there is a significant adaptive relationship

between ecology and morphology in North American and Southern African flycatcher

communities. Second, using morphological traits and observations on foraging behavior,

I test whether ecomorphological relationships are dependent upon locality. Third, using

multivariate discrimination and cluster analysis on a morphological data set of five

flycatcher clades, I address whether there is broad scale ecomorphological convergence

among flycatcher clades and if morphology predicts a course measure of habitat

preference. Finally, I test whether there is a common morphological axis of

diversification and whether relative age of origin corresponds to the morphological

variation exhibited by elaenia and tody-tyrant lineages. The general results were this: 1)

Morphology significantly predicted the foraging behavior in both NA and SA flycatchers.

2) North American and Southern African communities are concordant with respect to the

ecomorphological relationships. 3) I found that there are lineage-specific positions in

morphological space when examining synoptic morphological samples of Old and New

World flycatcher clades. 4) There were fundamental differences in the orientations of

morphological disparity of Old versus New world flycatchers. 5) The results of separate

principal components and common principal component analyses reveal a larger morphological volume being occupied by older lineages, but 6) Macroevolutionary patterns exist within constituent clades that are inconsistent with a Brownian motion evolutionary hypothesis. Hence, the differences in morphological disparity observed in flycatchers are most likely due group-specific factors including genetic constraints, ecological limiting similarity and ecological opportunity. Phylogenetic constraints to morphological evolution are not universal throughout the history of flycatcher evolution.

Furthermore, historical and/or stochastic factors were equal to ecological factors during flycatcher evolution.

Approved: Donald B. Miles

Associate Professor of Biological Sciences

5 ACKNOWLEDGEMENTS

Foremost, I thank my wife Nancy. Her love, patience, and encouragement have been there from the start and continue to this day. My son Charlie (10 months old at the time of this writing) has added lots of encouragement for this project in a special way.

Opie, as always, helped me keep things in perspective. These members of my family, above all, made my experiences at Ohio University great ones and I am dearly indebted to them.

I want to thank my parents, Nancy’s parents and my grandmothers for lots of love, encouragement and financial support during this dissertation project. I could not have done it without them. Also, my siblings were there for me when I needed them and

I appreciate their support. I especially want to thank Rocky for his time and encouragement on many occasions. His perspectives got me through a lot of tough times during research and writing. I want to thank my extended family in general for their support and high expectations.

Don Miles gets a lot of credit for this project. Since my master’s thesis, he has influenced my worldview in positive ways and taught me to look at the ecology of organisms, especially birds, from an evolutionary perspective. John Scheibe also gets a lot of credit for pointing me in the right direction when it comes to comparative biology.

These men taught me the importance legacy and chance current ecological communities.

This project has been improved greatly by comments and suggestions from my proposal and dissertation committee members – Audrone Biknevicius, Mark Dybdahl,

Jim Dyer, Molly Morris, Steve Reilly and Willem Roosenburg. I also want to thank the

6 Ohio University Evomorph group for helpful comments. Members from the laboratories of Audrone Biknevicius, Molly Morris, Willem Roosenburg, Larry Witmer and J. Van

Remsen Jr. have also lent support and helpful comments and suggestions.

Thanks to T. Smith and two anonymous reviewers of the “elatody” manuscript

(Chapter 4) for helpful comments and suggestions. Also, I would like to thank the natural history museums and staff of the Transvaal Museum (T. Cassidy), Louisiana State

University (J. V. Remsen Jr.), University of Kansas (M. Robbins), Field Museum (D.

Willard), and National Museum of Natural History (J. Dean) for museum loans and access to their collections. Thanks to Kevin de Queiroz, Mark Lensmeyer and Shaun

Pharr for putting me up (putting up with me) and feeding me while collecting museum data.

For help while in the field, I thank Stacey McKinnen, Sara Kozup, Dave

Rushworth, Alan Kemp, Robert Ricklefs and the Transvaal Museum staff. I thank Laurie

Dries, Scott Moody, Jim Robbins, John Scheibe, Jayc Sedlmayr and Martin Wikelski for helpful discussions about the research. Pete Larson and Chris Klingenberg helped considerably with common principal components and bootstrapping programs. Finally, thanks to my office mates: Rick Essner, Lance McBrayer for helpful comments on my research and appropriate ribbing.

7 TABLE OF CONTENTS

Abstract...... 3

Acknowledgements...... 5

Table of Contents...... 7

List of Tables ...... 9

List of Figures...... 12

List of Appendices ...... 15

Chapter 1. Introduction to the Dissertation...... 16

Background and Interests ...... 17

Chapter Summaries ...... 20

Chapter 2. Concordance among North American and Southern African Flycatcher

Communities...... 24

Introduction...... 25

Methods...... 28

Results...... 38

Discussion...... 43

Chapter 3. Convergence in the Adaptive Radiations of Old and New World

Flycatchers ...... 51

Introduction...... 52

8 Methods...... 55

Results...... 61

Discussion...... 70

Chapter 4. Comparative Morphological Disparity of Elaenia and Tody-tyrant flycatcher

Clades ...... 77

Introduction...... 78

Methods...... 80

Results...... 87

Discussion...... 92

Chapter 5. General Conclusions and Future Directions...... 98

Literature Cited ...... 101

Tables...... 121

Figures...... 139

Appendix I ...... 164

Appendix II ...... 192

Appendix III...... 208

Appendix IV...... 214

9 LIST OF TABLES

Table

2.1 Cross tabulated data entered into the correspondence analysis on search behavior.....

...... 121

2.2 Coefficients of eigenvectors, eigenvalues and cumulative explained variance for

correspondence analysis on foraging location in New World flycatchers...... 122

2.3 Foraging percentages for foraging mode used in correspondence analysis of attack

behavior of Old and New World Flycatchers ...... 123

2.4 Coefficients of eigenvectors, eigenvalues and the cumulative explained variance for

correspondence analysis on attack mode of foraging ...... 124

2.5 Morphological measurements of flycatcher species used in test of ecomorphological

concordance ...... 125

2.6 Coefficients, eigenvalues and cumulative explained variance for the principal

components analysis of morphology of Old and New World flycatchers...... 126

2.7 Summary of canonical correlation analysis on search behavior...... 126

2.8 Summary of canonical structure of morphology and search behavior in New World

flycatchers...... 127

2.9 Summary of the canonical correlation analysis on morphology and attack behavior

in Old and New World flycatchers ...... 127

10 List of Tables. Continued.

2.10 Summary of canonical structure of morphology and attack behavior in Old and New

World flycatchers...... 128

2.11 Results of ANCOVA testing for differences in locality means in the foraging with

morphological covariates...... 128

2.12 Attack rates and standard errors based upon grand averages of species with n

individuals drawn from the total data set...... 129

3.1 Coefficients of the principal components analysis on separate covariance matrices .....

...... 130

3.2 Descriptive statistics of the principal components analyses on separate covariance

matrices...... 131

3.3 Results of canonical discriminant analysis on nine morphological variables of Old

and New World flycatchers ...... 132

3.4 Lineage centroids for the canonical discriminant analysis ...... 132

3.5 Squared Mahalanobis distances and F statistics among lineages of Old and New

World flycatchers...... 133

3.6 Misclassification table of Old and New World flycatcher lineages based upon the

nine morphological variables and a prior classification rate proportional to the

sample size of the group ...... 133

3.7 Statistical moments of the ellipse analysis...... 134

11 List of Tables. Continued.

3.8 T-values of pair wise comparisons in orientations and shape of ellipse parameters

among lineages...... 135

4.1 Coefficients and cumulative variance explained in the 1) separate principal

component analyses (PCA) and 2) common principal component analysis (CPCA) on

elaenia and tody-tyrant lineages...... 136

4.2 Estimated t values of sequential pair-wise comparisons between subclade parameters

of ellipse shape and orientation...... 137

4.3 Median Nearest Neighbor Distances (NNDs) for each lineage and subclade ...... 138

12 LIST OF FIGURES

Figure

1.1 A phylogeny of the Passeriformes based upon Sibley and Alquist (1990) DNA

Hybridization data...... 139

2.1 The phylogeny of the flycatcher species used in the study of concordance based

Sibley and Alquist (1990) DNA Hybridization data...... 140

2.2 Joint scatterplot of the species scores in search foraging of New World flycatchers.....

...... 141

2.3 Joint scatterplot of the species scores in attack foraging of Old and New World

flycatchers...... 142

2.4 a & b. Scatterplot of the average species scores of Old and New World flycatchers

from a principal components analysis on log10 transformed morphological variables...

...... 143

2.5 Scatterplot predicting search behavior from morphology in the New World

flycatchers...... 144

2.6 a & b. Scatterplot of the canonical correlation analysis of attack behavior and

morphology...... 145

2.7 Attack rate plotted against the first morphological PC...... 146

2.8 Percent of foraging maneuvers of Old and New World species that are comprised of

flycatching...... 147

13 List of Figures. Continued.

3.1 Scatterplot of the first and second PC axes from separate covariance matrices of three

flycatcher lineages ...... 148

3.2 Frequency distributions of species scores along the first three principal components

for each group ...... 149

3.3 Sign, magnitude and similarity of each variable within each eigenvector of the

separate principal components analyses of three flycatcher lineages...... 150

3.4 The relative rates of overall body size and shape evolution ...... 151

3.5 The morphological space occupied by the five lineages of flycatchers in canonical

discriminant space...... 152

3.6 The morphological space occupied by the five lineages of flycatchers in canonical

discriminant space...... 153

3.7 UPGMA cluster analysis on size-free generic mean morphological traits...... 154

3.8 The evolutionary rates and changes in tempo of diversification in flycatcher

subclades...... 155

4.1 Lineage specific patterns of diversification in bivariate space ...... 156

4.2 Synthesized phylogeny from Sibley and Alquist (1990) DNA and Lanyon (1988a;

1988b) morphological data ...... 157

14 List of Figures. Continued.

4.3 Standard errors about bootstrapped coefficients plotted against the eigenvectors of a

common principal components analysis on six log-transformed morphological traits

of elaenia and tody-tyrant flycatcher lineages ...... 158

4.4 Scatterplot of elaenia along the first two common principal component axes ...... 159

4.5 Scatterplot of tody-tyrants along the first two common principal component axes .160

4.6 Scatterplot of elaenia along the first second and third common principal component

axes ...... 161

4.7 Scatterplot of tody-tyrants along the second and third common principal component

axes ...... 162

4.8 Total lineage morphological disparity in of elaenia and tody-tyrant flycatchers based

on a characterization of the morphological volume from a common principal

components analysis ...... 163

15 LIST OF APPENDICES

Appendix

I Species, sample sizes and morphological means and standard deviations for tyrant

flycatchers...... 164

II Species and morphological data used in the CDF, UPGMA and ecological axis

analyses...... 192

III Species used in the CPC and Lineage disparity analyses, lineage membership

designations, number of specimens measured and mean morphological

measurements by species for each variable ...... 208

IV Results of the multivariate ellipse analysis on the CPC scores of Elaenia and Tody-

tyrant lineages, and all included subclades...... 214

16

CHAPTER 1. INTRODUCTION

Clay E. Corbin

17 The purpose of this introduction is to give the reader an overview of my background leading up to my dissertation interests and also to give an outline of the science I have attempted to address in each chapter. Each research section of the dissertation will be summarized by a chapter-specific abstract.

BACKGROUND AND INTERESTS

My master’s thesis (Corbin 1995) was centered on the hypothesis that

interspecific competition structures avian communities. Species that utilize the

environment in very similar ways (sensu Root 1967) should not be able to coexist (Gause

1934; MacArthur and Levins 1967). I tested this idea using the emergent qualities

(species richness and diversity) and the concept of multivariate niche breadth and overlap

(Hutchinson 1959; Harner and Whitmore 1977). I inferred each community member’s

distribution in ecological space from morphological measurements on each species – a

method central to Ecological Morphology (Bock and von Wahlert 1965; Karr and James

1972). Then I compared the observed niche metrics for communities of various sizes

against those generated from random assortments of bird species into “null” communities

over the same range of species richness. If competition were structuring the observed

communities, one would expect smaller niche breadths and niche overlaps than the

randomly generated communities. The species pool from which I randomly constructed

the null communities was comprised of all bird species on the research site, regardless of

guild membership or phylogenetic affinity. As a result, I found little evidence that

competition was determining the make-up of the communities. Indeed, if competition is

an important structuring mechanism to bird communities, it would probably be observed

among those species that are utilizing the environment in the most similar way instead of

18 throughout the entire local species pool. Hence, I turned my interests to guild membership for testing competition – primarily between active foragers (i.e. warblers and vireos) and sit-and-wait foragers (the flycatchers). Again, I found little evidence that the

“ghost of competition past” has had a guild-wide effect on the morphologies of these species (Corbin unpublished). Nonetheless, my interests in the sit-and-wait foraging mode and the diversity of the world’s flycatching birds were initiated.

There are a few things, pertinent here, that one learns while studying North

American flycatchers. First, North America is relatively depauperate of tyrant flycatchers. A survey of the South American Tyrannidae (Traylor and Fitzpatirck 1982) reveals that in some habitats, one in every four passerine species is a tyrant flycatcher – not a trivial statistic when referring to Neotropical avian biodiversity. Second, even though flycatching – the behavior of waiting for and then attacking a passing prey item – is a common foraging mode to most of the 415 species of Tyrannidae, there is variation in foraging mode. Foraging mode varies from classic flycatching (i.e. Contopus spp.) to gleaning (Elaenia spp.) in open habitats (i.e. Tyrannus spp.) and dense vegetation (i.e.

Myiarchus spp.). Third, sit-and-wait foraging is not restricted to the New World.

Flycatching is a foraging mode that has been repeated along several independent lines of passerine birds. Examples across the globe include the tyrant flycatchers (Tyrannidae) to three tribes in the Corvidae (the Fantails (Rhipidurini), Drongos (Dicrurini) and Monarch

Flycatchers (Monarchini)) to a tribe in the Bombycillidae – the Silky flycatchers

(Ptilogonatini) and finally the “Old World Flycatchers in the family Muscicapidae (Tribe

Muscicapini). For this dissertation, I concentrate on the evolutionary and

19 ecomorphological patterns in three of the largest clades of flycatchers: the Tyrannidae, the Monarchini and the Muscicapini.

To put these groups in a phylogenetic perspective, the monarch and muscicapine flycatchers (Old World Flycatchers) are more closely related to each other than either is to the tyrants. The Old World groups are included in the larger clade of oscine passerine birds and the tyrants are one of the large clades within the sub-oscine passerine birds

(Figure 1.1). When an ecologist or evolutionary biologist observes a repeated and seemingly adaptive phenomenon such as convergence in sit-and-wait behavior, they may ask “how tightly matched is the convergence?” In other words, have the independent ecological histories of flycatching clades been the same? If different, do these differences explain differences in the total ecomorphological variation that is seen among the clades today?

I approach this problem on three levels (chapters 2-4). First, at the community level, I address whether the phenotypic variation among different flycatcher communities

(North America and Southern Africa) is similar and whether it has responded in a similar way to their respective environmental milieus. Second, I expand the idea of concordance among communities to test for differences among entire flycatcher clades. And finally, I ask whether the similar biogeoraphical histories of two clades within the New World flycatchers have produced similarities in morphological variation.

20 CHAPTER SUMMARIES

Chapter 2 – Community Concordance in North American and Southern African

Flycatchers

In both the Old and New Worlds, independent clades of sit-and-wait insectivorous birds have evolved. These independent radiations provide an excellent opportunity to test

whether the relationships between morphology and ecology are concordant among

different communities of flycatchers. First, with canonical correlation analysis, I test

whether there is a significant relationship between ecology and morphology in North

American (NA) and Southern African (SA) flycatcher communities. Second, using

multivariate characterizations (Principal components and reciprocal averaging) of

morphological traits and foraging behavior, I test, using ANCOVA whether the

ecomorphological relationship is dependent upon locality after taking morphological

differences into account. Also, I accounted for phylogenetic relationships among the

species in the two communities to see if observed patters were consistent with an

adaptive explanation of the form-function relationships. I found that morphology predicted the foraging behavior in both NA and SA flycatchers. Also, the results of the

ANCOVA showed that these communities are concordant with respect to the ecomorphological relationships. Also, the phylogenetic relationships among the species did not seem to influence the patterns seen in these data. Hence the ecomorphological relationships are adaptive and common to both communities of flycatchers. However, the relationships between ecology and morphology (explained variances) are not as tight as

21 one may predict. This may be due to differences in habitat or broad scale historical influences between NA and SA flycatchers.

Chapter 3 – Morphological Evolution and Niche Partitioning in Old and New World

Flycatcher Clades

Adaptive radiations are exciting phenomena in evolutionary ecology. Except for a few cases, the concept of evolutionary convergence has not been applied to lineages that have undergone adaptive radiation. The independent lineages of Old and New World flycatchers provide an excellent opportunity to test for generalities in large scale, independent macroevolutionary events. Using multivariate and cluster analysis on a morphological data set of five flycatcher lineages, I address questions to shed light on convergent adaptive radiation. First, is there broad scale ecomorphological convergence among flycatcher lineages? Second, does morphology predict habitat preference across flycatcher genera regardless of phylogenetic affinity? I found that there are lineage- dependent positions in morphological space. Also, there were fundamental differences in the orientations of morphological disparity of Old versus New world flycatchers.

However, the cluster analysis revealed that there is some level of matching across genera away from what one would predict according to phylogenetic or geographic nearest neighbors. Clusters predict the habitat preference across many of the genera under consideration. At the level of species, there is little evidence of convergence and modest evidence for phylogenetic conservatism. However, at the level of genera, convergence is apparent and seems to be associated with preference for open or closed habitats. Hence, there may be more than one way to adapt to an ecological problem but at fine scales morphological evolution may be contingent upon history and phylogeny.

22 Chapter 4 – Morphological Disparity and Diversification in Elaenia and Tody-Tyrant

Flycatcher Clades

We compared the morphological disparity of two tyrant flycatcher lineages, the elaenia and tody-tyrants. These species-rich lineages are hypothesized to have undergone similar evolutionary radiations. However, these lineages differ in age of origin suggesting that the older tody-tyrant lineage should be more morphologically diverse than the elaenia. In this study we test whether the relative age of origin corresponds to the morphological disparity exhibited by these lineages. We compared the morphological volumes of these lineages using principal components analyses of bill, wing, tail and leg characters.

Results of separate principal components and common principal component analyses reveal a larger morphological volume being occupied by the older Tody-tyrant lineage.

Also the tody-tyrants seem to have diversified to a larger extent in size-free morphological volume. Further, we analyzed the historical trends throughout the evolution of these lineages by examining the size, shape and orientations of 95% confidence ellipses around nested monophyletic groupings. This revealed a general lineage-wide maintenance of ellipse statistics when size is included. Hence, changes in body size played a major role in the early evolution of both of these lineages. However, when analyzing the trends in shape, these similarities breakdown. Both lineages exhibit independent evolutionary trajectories. Furthermore, there was not a linear relationship between age of origin and morphological disparity. Macroevolutionary patterns exist in particular groups that are inconsistent with a Brownian motion evolutionary hypothesis.

Both lineages contain monophyletic subclades that are either under- or hyper-dispersed in their respective morphological volumes. We conclude by hypothesizing that the

23 differences in morphological disparity observed in these groups of flycatchers are possibly due group-specific factors including genetic constraints, ecological limiting similarity and ecological opportunity. Unique historical or stochastic factors probably played an equal role as ecological factors in the morphological evolution of these flycatcher lineages.

24

CHAPTER 2. COMMUNITY CONCORDANCE IN NORTH AMERICAN AND

SOUTHERN AFRICAN FLYCATCHERS

Clay E. Corbin

25 INTRODUCTION

Independent clades of sit-and-wait insectivorous birds have evolved several times.

Three of these radiations have been extensive – the Tyrannidae in the Neotropics and the

Monarchini and Muscicapini of Old World biogeographical realms. While it appears the

Tyrannidae had an Old World origin, their radiation into one of the largest passerine

groups occurred primarily during the isolation of South America (Keast 1972; Traylor

and Fitzpatrick 1982). Separate communities of these birds interact with their environment in similar ways. Hence, the existence of flycatcher communities provides an excellent opportunity to test whether the relationship between phenotypic and ecological variation is similar among localities – a test of ecomorphological concordance

(Miles and Ricklefs 1984; Schluter 1986; Miles et al. 1987; Losos and Miles 1994;

Lamouroux et al. 2002; Miles, Corbin and Pearson in review). Is there a common pattern

of interaction between the phenotype and ecology among non-related bird communities?

Ecomorphological analysis is concerned with inferring ecological relationships of

organisms from morphological traits (Leisler and Winkler 1985, 1990; Bock, 1990, 1994;

Karr and James 1975; Ricklefs and Travis 1980; Miles and Ricklefs 1984; Ricklefs and

Miles 1994; Wainwright and Reilly 1994). This type of analysis is especially interesting

here because there are distant phylogenetic relationships between Old and New World

flycatchers. If the species in North America and Southern Africa are concordant with

respect to the functional relationships between the phenotype and their environment, this

provides evidence for repeated phenotypic adaptation (Coddington 1988). Hence, is the

relationship between ecology and morphology in each locality a reflection of local

26 adaptation or of recent common ancestry? Furthermore, are these relationships similar among Old and New World flycatcher communities when accounting for phylogenetic information?

Similarities among flycatcher communities in terms of foraging behavior and morphology have been superficially addressed in a few investigations (Keast 1972, Karr and James 1975). While strikingly similar in some cases (see Karr and James, 1975) these clades have had quite different biogeographical and ecological histories, including differential levels of potential competition (Keast, 1972). Ecomorphological concordance among localities may be expected when phenotypic change is adaptive and the same ecological factors are common to the assemblages. On the other hand, the differences in an ecomorphological relationship among communities of birds can reflect historical differences, or environmental differences.

While some studies were successful in predicting relative sit-and-wait foraging from morphology (lizards, Pianka 1973; Scheibe 1987; birds of prey, Jaksic and

Carothers 1985; Gamhauf et al. 1998; passerines, Keast et al. 1995), others have cast doubt on the idea that this may be common to all systems (Weins and Rotenberry 1980;

Motta 1988). Typical foraging maneuver called “flycatching” or “hawking” of many Old and New World flycatchers involves a bird waiting on a perch for a prey item to appear, launching to capture the prey in its bill and then returning to a perch to consume the prey and resume scanning. However, even though flycatching is typical and maneuvers such as pecking or probing are rare (Traylor and Fitzpatrick 1982; Fitzpatrick 1980; 1985), members from both Old and New World flycatcher groups lay along a continuum of relative specialization in sit-and-wait foraging. For example, in the New World

27 Tyrannidae, Serpophaga species utilize an active foraging mode that includes many gleaning maneuvers and relatively few flycatching maneuvers while Contopus species primarily exhibit sit-and-wait flycatching (Fitzpatrick 1980). In the Old World

Muscicapidae, Myioparus and Melaenornis species are respectively different (Fraser

1983). In addition, there is variance in the habitat preferences among flycatcher species.

Tyrannus and Maleanornis species utilize open habitats while many Empidonax and

Terpsiphone species are found foraging in closed vegetation (pers. obs.). Hence, in both communities there is a range of habitat preference and foraging behavior that is potentially concordant and predictable by morphology.

Some ecomorphological relationships have been established within Neotropical flycatchers (Fitzpatrick 1980; Sherry 1984) and in comparison to other species in

Neotropical (Miles et al. 1987) and North American communities (Cody and Mooney

1978; Eckhardt 1979; Miles and Ricklefs 1984). Also, some work has been done on the adaptive external morphology of Old World flycatchers (Erard 1987; Korzoun et al.

2000) and, in general, the patterns seen in New and Old World flycatchers correspond to what one would predict from past literature on ecomorphology in the Passeriformes. The bill is used for handling and ingesting food. As a result, the bill is reliable as a predictor of ecology (Beecher 1951; Bock and von Wahlert 1965; Schoener 1965; Grant 1967;

Hespenheide 1973; Karr and James 1975; Miles and Ricklefs 1984; Leisler and Winkler

1985; Miles et al. 1987; Hertel 1994, 1995; Price et al. 2000; Forstmeier and Keller 2001; and many others). Tyrannid flycatchers that “spoon” prey items from the undersides of leaves tend to have long and wide bills while pincerlike long but narrow bills are exhibited by birds that pick prey from a substrate. Also, birds that utilize an aerial

28 hawking or “true flycatching” (Storer 1971) foraging mode tend to have relatively wide bills than those picking prey off the ground or from a substrate (Fitzpatrick 1985). Other morphological traits belie the natural history of a species. Wing and tail lengths indicate relative maneuverability (Norberg 1979; 1990) and hence habitat preferences among species (Leisler and Winkler 1985; Forstmeier et al. 2001). Long propulsive hand wings are found in species that use open habitats while short maneuverable wings are found in closed habitats (Fitzpatrick 1985). The length of the lower leg is a good predictor of habitat use and relative ground use (Carrascal et al. 1990; Block et al. 1991).

In this study, I use measurements on external morphology and observations on foraging behavior, and habitat from flycatcher species from Ohio, Arizona and southern

Africa to address two questions. First, does morphology predict foraging behavior within

Old and New World flycatcher communities? Second, are the communities concordant with respect to the ecomorphological relationships?

METHODS

Species and Study Areas

There are around 32 breeding sit-and-wait tyrannine bird species in North

America and 14 breeding sit-and-wait monarchine or muscicapine species in Southern

Africa (sensu Sibley and Alquist 1990). I was able to observe regional subsets of these

birds in Ohio, Arizona and South Africa. Observations for the African birds were

combined with published foraging data on those species (Fraser 1983) to increase the

number of species and sample sizes. The species used in this study are presented in

Figure 2.1.

29 Because there is some variance in flycatcher species with respect to the relative proportion of sit-and-wait to active foraging, I wanted to investigate the foraging behavior in comparison to species that are know to be active insectivorous foragers

(MaClean 1993; Barber et al. 2000, Jablonski 2001). Hence, I included a warbler species, the Painted Redstart (Myioborus pictus) and a combined sample of shrike species in the genus Batis (B. molitor, B. pririt). The batises were combined for two reasons.

First, in the field I could not always reliably distinguish the pririt batis (B. pririt) from the chinspot batis (B. molitor). Second, 2 x 4 and a 2 x 2 contingency analyses of Fraser’s

(1983) data accept a null hypothesis of no dependence in feeding technique (see below) and search location (in- or outside canopy) with respect to B. pririt and B. molitor (X2 =

3.3, df = 2, P = 0.19, and X2 = 3.7, df = 1 P = 0.054). Hence, there is little or no

statistical difference in how the batises are foraging that would affect this study.

Foraging Data Collection

Investigations were carried out primarily in the mornings (0500-1000) and

evenings (after 1600) in Ohio and Arizona May-August 1999 and 2001. The southern

Africa foraging data were collected also in the mornings (0800-1000) and evenings

(1400-1700) by myself during August of 1998 and by Fraser from 1979-1981 (Fraser

1983; Fraser pers. comm.).

Field sites were chosen to maximize habitat diversity and hence, species diversity

within the sit-and-wait insectivorous communities. Sites in Ohio included the Ridges

Landlab of Ohio University, Waterloo Wildlife Area, Deep Woods Farm (Ohio

Biological Survey), Stages Pond State Nature Preserve, Wahkeena Nature Preserve and

Rising Park (Lancaster). Sites in Arizona were Cave Creek Canyon, Rustler and Barfoot

30 Park areas in the Chiricahua Mountains, Ramsey Canyon in the Huachuca Mountains,

Madera Canyon in the Santa Rita Mountains, Patagonia-Sonoita Creek Preserve,

Patagonia Lake State Park and Las Cienegas National Conservation Area. South African sites included Nylsvley Nature Reserve, Suikerbosrand Nature Reserve, the southern region of Kruger National Park, Bophuthatswana (North of Pretoria), the Rushworth ranch near Hoedspruit, and the False Bay, Hluhluwe, Kranzkloof and Stainbank Nature

Reserves of KwaZulu-Natal.

I collected sequential foraging observations on target species listed in Figure 2.1

(Altmann 1974; Remsen and Robinson 1990). I attempted to choose variables that reflected both the searching and attacking aspects of foraging (Remsen and Robinson

1990). The following information reflects searching behavior: perch height (PH)

(meters), height of perch structure (TH) (meters), and the distance to the nearest vegetation above the bird (OC). Because there were many instances where a bird was not under any cover at all (typical of the Tyrannus spp.), this latter variable was categorized as either under structure (“C” for closed) or open (“O”). Note that the open category includes wide-open (i.e. grassland), open forest gaps, or birds foraging at the tops of trees

(i.e. Contopus pertinax). Perch height (PH) was categorized to be consistent with Fraser

(1983) as follows: PH1 = < 1 m, PH2 = 1-2 m, PH3 = 2-3 m, PH4 = 3-4 m, PH5 = > 4 m.

An index to relative bird perch position with respect to the vegetation height was calculated as (RPH = PH/TH). The attack aspect of foraging was simply the species foraging mode. This ranged from 1) flycatching – which I defined as an airborne bird attempting to capture an airborne prey item, 2) hover-gleaning – airborne bird with stationary prey item, 3) standpicking – maneuver where no flight by the bird is required

31 and the prey item is stationary and, 4) pouncing – the bird flew to the ground for either stationary or mobile prey.

Sequential observations allowed the calculation of an interspecific measure of foraging performance – that of attack rate (AR). This is the mean number of times per second a bird attempts to capture a prey item. Hence this measures the relative amount of active foraging of a species (Cody 1974; Eckhardt 1979; Robinson and Holmes 1982;

Remsen and Robinson 1990) and provides a quantitative link between morphology and behavior (Arnold 1983; Losos 1990). Search time was recorded by stopwatch as the amount of time a bird searched for a prey item to the time it attempted to capture an item.

Handling time was negligible in most cases so when the bird landed the time recording began. If there was a noteworthy handling time, it was noted in the field and the observation was excluded from analysis. Because there was variance in the number of times any one bird was observed attempting to capture a prey item, I selected individuals from each species for which there was a series of three sequential observations (the first and next two observations). Then, I averaged the search time for the series. Finally, I computed a grand average for each species. Therefore, the attack rate for a species is estimated by the number of attempts at a prey item per second.

The non-independent nature of sequential observations may bias statistical results

(Morrison 1984; Hejl et al.1990, Leger and Didrichsons 1994; Jenkins 2002). Variation in foraging behavior tends to stabilize when there are at least 30 independent observations (Morrison 1984). For this study, the total number of individuals was less than that for some species. Therefore, I assessed whether using all (sequential) or just the first (independent) observations for each species would influence the interspecific

32 relationships among species in multivariate foraging space using ANOSIM (Primer-5

2001). I computed the median and mean values for categories of TH, RPH, OC and FM, using all observations and then using only the first observations on each species. Two

Bray-Curtis similarity matrices (one using the average and one the median values) of the species based on the foraging variables were generated using dataset (= “all obs” or “first obs”) as the independent factor. I compared the observed similarity matrices to 1000 comparisons of randomly permuted similarity matrices (Primer-5 2001) to test for similarity between the full and first data sets.

Morphological Data Collection

The natural history museums of the University of Kansas in Lawrence, the Field

Museum in Chicago, the Transvaal Museum in Pretoria, Ohio University in Athens and

Louisiana State University in Baton Rouge provided skins for measurement.

Morphological characters were chosen due to their likely representation of adaptations to different aspects of the environment (Leisler and Winkler 1985). I measured total length, wing length, and tail length with a ruler (nearest 0.5 mm), and bill depth, length (tip to naso-frontal hinge, width 1 (across quadrates) width 2 (across nares), tarsus and middle toe with dial calipers (nearest 0.1 mm). All morphological data were log10 transformed

prior to analysis. Sample sizes ranged from 1 to 30 individuals per species.

Phylogeny

The distance tree of Sibley and Alquist (1990) provides the best estimate of a

phylogeny comprised of all the target species. In addition, one can use the distance

values to infer evolutionary branch lengths. The values in the tree are derived from

differences in melting temperatures between duplex combinations of inter and intra-

33 specific DNA molecules (Sibley and Alquist 1990). The smaller the value, the more closely related two taxa are. For the species in this study the values range between 19.4 and 2.4 (change in value = 1.0 ~ 4.7 million years Sibley and Alquist (1990)). The reduced tree for the species used in this study contained several nodes near the tips with no branch length information. I arbitrarily set branch lengths above those nodes to 0.1 for congeners (Tyrannus, Myiarchus, Sayornis, Contopus and Empidonax) and to 0.5 for non-congeners (Melaenornis pammelina, Bradornis mariquensis and Sigelus silens;

Muscicapa striata and Myioparus plumbeus) (see Figure 2.1).

Because the proximity of species in ecomorphological space may be affected by common ancestry and not because of independent coevolutionary events, I computed phylogenetically independent contrasts (Felsenstein 1985, Harvey and Pagel 1991; Rohlf

2001) using CAIC (Purvis and Rambaut 1995) of the averaged species scores from the principal component and principal coordinate analyses. I assumed the polytomies were soft (Purvis and Garland 1993). For comparison between relationships with and without phylogenetic information, I used both the contrasts and the log10 transformed variables in

the ordination and regression analyses (see “Statistical Analyses”). I ran regressions

through the origin for the contrast comparisons (see Purvis and Rambaut, 1995).

Statistical Analyses

Foraging behavior

I performed separate correspondence analyses (CA) using the statistical package

MVSP (2000) on search and attack aspects of the foraging observations. CA is a dual ordination on the rows and columns of a contingency table where the cells are enumerated categorical values (MVSP 2000). Here, the values are the percentages of

34 cross-classified search locations or attack mode percentages used by each Old and New

World flycatcher species. CA axes that are extracted from those tables successively explain smaller proportions of the column and row variance. I used the scores along the first three CA axes in subsequent analyses. Also, independent contrasts were computed on the scores for comparison with original data in both search and attack space. The table for the attack behavior consisted of species along the rows and percentages of FM along the columns. For the search aspects, I cross-classified PH and OC and computed the percentages for each cell in the resulting matrix. Only the New World species were included in the investigation of morphology and search location because Fraser (1983) presents only separate percentages for PH and OC. This renders cross-classification impossible.

Morphology

I used principal components analysis to reduce the morphological data set to a few uncorrelated new variables that explain a majority of the available variance in the original data. Principal components analysis (PCA) is similar to correspondence analysis but here, the data are continuous measurements of morphological traits instead of enumerated percentages. The swarm of morphological data in Euclidean space is rotated so that the axis of greatest variance is the first PC. The directional cosines (eigenvectors) are computed for the first and subsequent orthogonal axes that explain successively smaller amounts of the variance. New variables (scores along the axes) were extracted from the covariance matrix of the log-transformed, morphological variables.

The unbalanced sample sizes for species (n = 1-30) may skew results of a PCA towards species that are better represented. To investigate if there was an effect of the

35 unbalanced morphological matrix, I compared the eigenvectors of the PCA where all individuals are included to the eigenvectors where only the species means for each variable is used. The results were the same. Hence, I averaged the scores for each species from the PCA on all individuals included for the analyses below.

The first PC axis usually reflects variation in overall body size. Because all PC axes are orthogonal to each other, I consider the remaining axes to be relatively free from body size (Burnaby 1966; Klingenberg in press). Finally, I computed independent contrasts on species scores for morphology in PCA space for comparison.

Tying morphology and behavior together

I investigated the correspondence of morphological and behavioral data using two separate canonical correlation analyses (CCA) and an analysis of covariance (ANCOVA)

(Miles et al. 1987). Because the ratio of morphological variables (nine) to number of species (k = 24) was large, I reduced the morphological data by using only the scores along the first three PC axes. The scores for these three axes were entered as the morphological data into the CCA. The behavioral variables of the two CCAs were species scores from the first three search CA and first three attack CA axes.

The canonical correlation analysis examines the relationship between a linear combination of the X variables (PCA scores) and a linear combination of a set of Y variables (search or attack CA scores) (SAS 2001). I computed correlation coefficients between the morphological combinations and their behavioral counterparts. The null hypothesis in a CCA is that the first canonical correlation and ones that follow were zero.

To test for an adaptive canonical correlation between behavior and morphology, I repeated the analysis using independent contrasts of PC and CA scores.

36 To test the hypothesis of concordance, the scores from first, second and third attack canonical axes were entered into an ANCOVA. An analysis of covariance has two main aspects. First, it tests for homogeneity of regression slopes among groupings and second, it tests for differences in means of a dependent variable after adjusting for covariates. The possible outcomes of an ANCOVA describe four different situations (see

Miles et al. 1987). First, there are no differences in either the slope or mean aspects.

Second, there are differences in both aspects. Third and fourth, there is a difference in only one aspect. In this study, the independent variable entered into the analysis was locality (Old or New World), and the species scores along the morphological axes were treated as covariates. Hence, the ANCOVA was used to 1) test that the Old and New

World ecomorphological regressions were similar and 2) test for homogeneity of attack

CA axes between Old and New World flycatchers while holding the morphological variability constant. A significant interaction term (PC1, 2 and 3 by locality) would reject the null hypothesis of similar slopes (Sokol and Rohlf 1995) and hence reject the concordance hypothesis (Miles et al. 1987). For the second test, a significant main effect after adjusting the means for the covariates would result from ecomorphological differences in locality based primarily upon morphological differences. Significant covariates identify which morphological variable(s) are significantly related to the response variables.

Phylogenetically independent contrasts using CAIC (Purvis and Rambaut 1995) are computed by summing weighted values where the weights sum to zero. At polytomies, the three or more taxa are divided into two groups based upon the tip value in relation to a weighted mean of all values (Purvis and Rambaut 1995). Then a contrast is

37 computed for the nodal value. Hence, the number of contrasts in a phylogeny with polytomies can actually be less than the number of species minus one. In this study, that loss in degrees of freedom resulted in a high variables-to-observations ratio (Stevens

1996) when using six variables (scores first three search and morphology axes) on the 12 contrasts in the CCA. An alternative was to use the same contrast data in a partial least squares (PLS) analysis (Klingenberg and Zaklan 2000; Rohlf and Corti 2000). Like

CCA, the method constructs pairs of variables from linear combinations of the original variables. However, the goal in pair construction is to maximize the covariation between the two sets of linear combinations. Hence, I entered contrasts of scores from search CA axes and morphology PCA axes into a PLS model using PROC PLS in SAS (2001). I extracted scores from the linear combination pairs and then computed correlation coefficients between them to investigate any loss of explanatory power in the prediction of search location from morphology when accounting for common descent.

It is necessary to run regressions on independent contrasts through the origin

(Garland et al. 1992; Purvis and Rambaut 1995). This quality of contrast data precludes their use in an ANCOVA because the model implicitly fits an intercept to regression lines when using class variables (SAS 2001) (but see Rohlf 2001). However I am interested in testing for convergence between independent clades with considerable phylogenetic distance among them (Sibley and Alquist 1990). Therefore, homogeneity between Old and New World ecomorphological relationships using the original data (rather than contrasts) is most likely a consequence of convergent adaptive response rather than statistical artifact (Coddington 1998).

38 Attack rate and morphology

In order to test whether morphology predicts a measure of interspecific foraging performance, I used linear regressions of attack rate against the first, second and third morphological PCs. There was no information on search time in Fraser’s observations so

I used only the species for which I had ample information.

RESULTS

Foraging

A total of 1769 foraging observations across the New and Old World species were

recorded. The sample sizes for data sets with all and first observations varied with

respect to species and variable. In the test to see if there were differences using “all” or

“first” observations, only 14 of 1000 R (Rho) statistics (metric of similarity (Primer-5

2001)) generated randomly from the data sets were higher than the observed R values

(global R = -0.056, P = 0.006). Also, I am using medians for the CA analyses; therefore,

the intraspecific variance that is added by using all observations would not obscure

interspecific patterns of foraging (Morrison 1984). Therefore, with respect to foraging in

these 23 species, I assume that the intraspecific variance is small compared to the

interspecific variance and hence, use the median values for pooled observations in

subsequent analyses (Leger and Didrichsons 1994; but see Jenkins (2002) for discussion

on instances where intraspecific foraging variability may be important).

39 Search Aspects

The cross-categorized data of PH and OC used in the search analysis are in Table

2.1. The first three axes of the CA on the search aspects of New World foraging flycatchers explain 43, 25 and 9% for a total of 77% of the total variance (Table 2.2).

The first axis is a contrast between open (O) and closed (C) habitats and the second axis is a continuous gradient of low to high perches (Figure 2.2). Species at the positive end of the first axis were usually found foraging within the forest canopy such as Empidonax virescens, Contopus virens, and Myiarchus tuberculifer. Birds in the open were the three

Tyrannus species and Contopus pertinax (at the tops of conifers). Birds searching at low perches (positive end of Search 2 in Figure 2.2) were Sayornis nigricans and Myiarchus cinerascens while, birds at high perches were Contopus pertinax and Tyrannus tyrannus.

Attack Aspects

Table 2.3 was used in the CA on attack foraging mode. The first three axes of the

CA on attack mode for all species explain 100% (57, 35 and 8% respectively) of the variance (Table 2.4). The first axis is a contrast between species that tend to pounce and standpick against species that hoverglean and flycatch (Figure 2.3). The second axis contrasts species that rely heavily on flycatching versus the other foraging modes. The third axis was a contrast between standpick and the other maneuvers. However, flycatching was relatively unimportant in explaining variance in the data in this axis as revealed by its near zero coefficient (Table 2.4). With the exception of the spotted flycatcher (Muscicapa striata), the Old World species are located in the extreme positions of the first two axes of attack foraging space. Myioparus plumbeus, the Batises and Terpsiphone viridis are primarily hover-gleaning, while the three closely related

40 muscicapine species Bradornis mariquensis, Maleanornis pammelina and Sigelus silens are pouncers. The New World species, on the other hand, are primarily flycatching.

Morphology

The species mean values for each variable and the sample sizes are presented in

Table 2.5. The coefficients that correspond to the eigenvectors of the PCA of species averages for morphological measurements are presented in Table 2.6. The corresponding species positions in morphological space are plotted in Figure 2.4a-b.

The first three axes of the PCA explain 93% of the total variance in the data. The first axis explains 79% of the variance and is considered a size axis because the coefficients are positive and roughly equal in magnitude. The second axis explains 9% of the variance and describes species with relatively large bill measurements at the positive end and large measurements for the other traits at the negative end. PC3 explains 4% of the variance in the data. In this axis, high loadings for tarsus length, toe length and bill width are at the negative end and high loadings for wing length, tail length and bill depth at the positive end of the axis.

Morphology and Search Behavior in the New World Flycatchers

Canonical correlation.—The first canonical correlation of morphology and search behavior in the New World flycatchers was significant (canonical R2 = 0.66, P = 0.03)

(Table 2.7). The first behavioral canonical axis is primarily associated with search CA2

(r = -0.94) and the first morphological canonical axis is associated with the first shape PC

axis 2 (r = 0.86) (Table 2.8). CA2 describes a searching gradient from low to high

perches. The morphological axis is a contrast between the bill measures at the positive

end and the other measures at the negative end. In Figure 2.5, the first morphological

41 axis for both the original PCA scores and their contrasts are plotted against the first search location axis.

The relationship between search behavior and morphology was maintained when accounting for phylogenetic information of species. While there was a drop in the explanatory power in the prediction of searching from morphology the first two pair of

PLS variates were significantly related to one another (intercepts at zero; R2 = 0.45, F =

8.9, P = 0.014, and R2 = 0.48, F = 10.486, P = 0.009 respectively), but the third pair were

unrelated to one another (R2 = 0.03, F = 0.3, P = 0.6). Plots of the first pair of PLS

variables are shown with the original data in Figure 2.5.

Morphology and Attack Behavior in the Old and New World Flycatchers

Canonical Correlation.—As in the search aspects of this study, only the first

canonical correlation was significant between morphology and attack behavior (r = 0.83)

(Table 2.9). In this analysis on both Old and New World species, the first canonical axis

of attack variables is primarily associated with CA1 (r = 0.74) (Table 2.10). This axis

contrasts species that tend to pounce and standpick against those that hoverglean and flycatch. The first axis of morphological variables is positively associated with the shape axis PC2 (bill traits vs. other traits) (r = 0.99). The first canonical correlation of independent contrasts is also significant (r = 0.89). However, while there is a significant canonical correlation, when taking phylogenetic information into account the factors important in explaining the behavioral variation shift from the second axis (compare

st roriginal = 0.99 to rcontrast = -0.15) to the first and third axes (1 , compare roriginal = -0.23 to

rd rcontrast = 0.68; 3 compare roriginal = 0.28 to rcontrast = -0.86). Nonetheless, when phylogeny

42 is taken into account morphology still predicts attack behavior. Hence, these patterns are consistent with an adaptive relationship between the ecology and morphology.

ANCOVA.—I used CA scores and PCA scores in an ANCOVA to test the hypothesis that the relationship between ecology and morphology between the communities was similar. The interaction terms for both foraging axes were not significant (attack axis 1: PC1 x Locality P = 0.31; PC2 x Locality, P = 0.93; PC3 x

Locality, P = 0.29, attack axis 2: PC1 x Locality P = 0.45; PC2 x Locality, P = 0.92; PC3 x Locality, P = 0.71). Hence, the initial assumption for the ANCOVA, that ecomorphological relationship between Old and New World flycatcher species were parallel, was satisfied and the interaction terms were subsequently dropped from the analysis. The resulting ANCOVAs are presented in Table 2.11. With respect to attack

CA axes 1 and 2, the ANCOVA models were significant (P = 0.0067 and P = 0.0003) and they explained 56% and 70% of the variance respectively. Unadjusted locality differences (Type I sum of squares; Table 2.11) were significant for both attack axes (P =

0.01, P < 0.0001 respectively). Hence, there appeared to be significant differences between the localities with respect to foraging behavior. However, after adjusting the means for the morphological covariates the significant differences vanished along the first attack axis (Type III sum of squares, Table 2.11, P = 0.29) but remained significant along the second axis (Type III sum of squares, Table 2.11, P < 0.001). An investigation of the covariates (Table 2.11) indicates the second morphological axis (bill shape vs. other morphological measurements) is related significantly to the first attack axis (pounce and standpick vs. flycatch and hoverglean; Type III sum of squares, P = 0.03). The first

(size) and third (tarsus length, toe length and bill width vs. wing length, tail length and

43 bill depth) morphological axes are significantly related to the second attack axis (flycatch versus other foraging modes).

Regression of Attack Rate and Morphology

Attack rate (Table 2.12) is negatively related to the first morphological axis (PC1) which is a size axis (Figure 2.7) (b = -0.68, F = 23.36, R2 = 0.59 P < 0.001 [contrast P =

0.009]). Batises and the Painted Redstart were not included in the regression analysis.

Small individuals at the negative end are associated with high attack rates, while larger

individuals such as Tyrannus species are associated with the positive end of the first axis

and subsequently have low rates of attack. Neither shape axis was associated with attack

rate in these species (PC2, R2 = 0.018, P = 0.59; PC3, R2= 0.1, P = 0.19).

DISCUSSION

The goals of this study were to determine if morphology predicts ecology within

Old and New World flycatcher communities and to test whether or not those

ecomorphological relationships were concordant between the localities. In the New

World flycatchers the search aspects of foraging measured in this study were predicted by

morphology. With respect to the attack aspects of foraging, both Old and New World

flycatchers attack aspects of foraging were also predicted by morphology. When I

accounted for phylogenetic information in both search and attack aspects, there was no

appreciable decline in the overall inference of behavior from morphology. Furthermore,

there appeared to be differences between localities with respect to their ecomorphological

relationships. When I adjusted the localities for differences in morphology, allometric

differences remained but the differences that were due to overall body size vanished.

Hence, with respect to the major predictor of foraging behavior in these flycatcher

44 species – body size – these localities are concordant. Also, as predicted there was a significant decline in attack rate with a decrease in overall body size but not with any aspect of phenotypic shape.

Searching Behavior and Morphology

The search location variable that explained a majority of the variance in the New

World flycatchers was whether or not species foraged in open areas. Perch height separated species in the second axis. There are two groupings of species with respect to the first axis (Figure 2.2) and the species in each grouping are what one would expect based upon the natural histories of these species (Ehrlich et al. 1998). Interestingly, the species foraging primarily in closed situations perch above 2 meters, while the species in more open environments are utilizing a full range of perch heights. These results may reflect the relationship between the area of the search image and habitat complexity observed in other flycatcher assemblages (Fitzpatrick 1981) and in other birds in general

(Moreno 1984; Corbin and Kirika 2002). In other words, if individuals are foraging in complex habitat where search area may be reduced in the horizontal plane, a species could compensate by foraging at a slightly higher elevation.

In ecomorphological space, the canonical correlation revealed that a significant correlation between perch elevation and morphological shape of the North American species. The relationship was maintained when phylogenetic information was excluded from the analysis. Hence, this pattern and the corroborating evidence of concordance between the localities are consistent with an adaptive hypothesis of the relationship between morphology and search position. Species with wide deep bills are foraging at higher elevations and species with relatively longer tails and leg measurements are

45 foraging near the ground. This makes sense functionally because relative leg length may impart an advantage with increasing ground use and wider, deeper bills may be advantageous to aerial foraging species in the open selecting larger prey items (Leisler and Winkler 1985). These results support the similar findings of Fitzpatrick (1985) and

Eckhardt (1979). However, neither study took into account phylogenetic information in their ecomorphological patterns and the “adaptive syndrome” (Eckhardt 1979) of flycatching was inferred from information at the tips of the phylogeny. Hence, this study lends support to the claims of these studies that within North American flycatcher communities, the link between search behavior and morphology has an adaptive underpinning. With respect to the Old World flycatcher community, more data are needed for basic natural history information (A. Kemp pers. comm. and W. Fraser pers. comm.) as well as to test that the relationship between search location and morphology also has an adaptive nature in the African flycatchers.

Attack Behavior, Rate and Morphology

The foraging mode that is important in explaining a large portion of the interspecific variance in the attack space is relative pouncing behavior. In the North

American species, phoebes (Sayornis) utilize this foraging mode the most. However, the proportion of pouncing maneuvers is small compared to South African Flycatchers. The muscicapine clade comprised of Southern Black, Fiscal and Marico flycatchers utilize this foraging mode to large extents when compared to the other foraging modes (see

Fraser 1983). Marico flycatchers utilize standpicking to a lesser extent and the other two species exhibit some flycatching. Here, the attack space is subdivided according to differences in foraging between Old and New World Flycatchers. The difference appears

46 to be based upon the amount of flycatching exhibited between the assemblages.

Examination of the attack foraging figure (Figure 2.3) reveals that there is a tight cluster of species associated with flycatching. All but one in that cluster are New World species.

The one Old World species is that of the muscicapine spotted flycatcher; a long distance migrant that winters in Southern Africa. Like the North American species, it primarily utilizes a flycatching foraging mode and only secondarily uses the other modes.

Interestingly, flycatching is of little importance in explaining the variance among these birds. This may be due to the different number of species that occur in each group coupled to the fact that it is the most common foraging mode in the North American species (Table 2.3 and Figure 2.8). It could be that pouncing behavior is simply an extension of the flycatching mode of foraging with a ground component (Fitzpatrick

1985). This is not a large component of foraging behavior in the North American species. Southern African muscicapine flycatchers may be utilizing this maneuver due to a ground foraging or pouncing ancestor. The muscicapines are closely related to species such as the saxicoline robins and chats, and both the muscicapines and saxicolines share a common ancestor with the turdine thrushes (Sibley and Alquist 1990). The species of these other groups utilize ground foraging (i.e. Catharus thrushes) and pouncing (i.e. bluebirds (Sialia)) maneuvers large extents (Power 1980; Wang and Moore 1990). An interesting way to examine the evolution of this behavior would be to map the percentages of pouncing utilization onto a phylogeny that includes representatives of muscicapine, saxicoline and turdine species (Brooks and McClennan 1991).

Morphology provides a good prediction of attack behavior in both the original and contrast data (Figure 2.6a & b). Although canonical correlation axes are a weak test of

47 the concordance hypothesis due to the common covariance matrix from which the group correlations are derived (Miles et al. 1987), the apparent differences in the elevation of slopes in Figure 2.6a were verified by the ANCOVA. If one were to control for morphological variation within the groups, there are differences in the means of the ecological scores. Given the highest correlations between the attack variables and their canonical variables are with CA1 (positively) and CA2 (negatively) (Table 2.10), the difference in elevation between the South African and North American flycatchers is due to the relative positions of the assemblages on those two axes. There is a slight shift to the right along the morphology axis of the Old World distribution of points with respect to the other flycatchers. This is also apparent in the contrast space. The shift hints of clade-specific (Lovette et al. 2001) patterns between Old and New World flycatchers that is also apparent in the ANCOVA. The largest contrast of the Old World (asterisk in

Figure 2.6b) is between the spotted flycatcher and lead-colored flycatcher. While these species are hypothesized to be phylogenetically closely related, there are large differences in both the morphology (Figures 2.4a & b) and foraging ecology (Figure 2.3). Hence, the short branch length at that node may not appropriately correspond to the amount of diversifying evolution that has taken place in the lineages leading to those species with respect to the rest of the relationships.

Attack rate decreases significantly with increasing size (Figure 2.7). This supports the findings from past work on search behavior in Neotropical flycatchers

(Fitzpatrick 1981; Sherry 1984) and other passerines related to the muscicapine flycatchers (Moreno 1984). Smaller birds are predicted be limited in the size of prey items they can effectively handle and consume (Schoener 1969; 1971 MacArthur 1972).

48 An exponential decrease in volume with decreasing surface area predicts that relatively larger numbers of smaller prey items should be taken to counter the increased energetic expenditure in smaller birds. This assumes that smaller prey items are more abundant than larger items. Future work should include the energetic expenditure of foraging movements as well as the relative abundance of both predator and prey in these flycatcher communities (see Sherry 1984; Bautista et al. 1998).

Ecomorphological Concordance and Differences among Foraging Aspects

The analysis of covariance reiterates what was found in the canonical correlation analysis with respect to the relationships between foraging behavior and morphology as well as the concordance between the flycatcher assemblages. After controlling for the morphological variance, there was no difference in CA1 distributions. However, in the

CA2 distributions (flycatching versus hover-gleaning) there was a significant locality effect. Hence, in attack space, with respect to the first axis, the ecomorphological trends are concordant due to the overlapping locality distributions along that first CA axis

(Figure 2.3). However, the locality based clumping along the second axis (Figure 2.3) leads to significant differences in the localities with respect to that axis.

In sit-and-wait foragers, the search location may be important to many aspects of the natural history of these species. That is because the perch may serve several different roles of foraging and vigilance for competitors and predators (Schoener 1971; Corbin and

Kirika 2002). The adaptive nature of the perch position is mirrored in this study. A comparison between Tables 2.7 and 2.9 reveal a difference in the relative fit of the different aspects of behavior on morphology. Specifically, morphology was a better predictor of attack foraging than it was of search location. In general, searching behavior

49 may be relatively unimportant in terms of energy expenditure to birds engaging in sit- and-wait foraging because of the multiple uses for a single perch. Hence, selection may be more important in the functional relationship between morphology and attack behavior rather than morphology and searching behavior. Corbin and Kirika (2002) demonstrated that sit-and-wait terrestrial foraging kingfishers (Halcyon albiventris) are opportunistic with respect to giving-up or attempting a prey attack from a given perch. Hence, search time or location may be relatively unimportant to a sit-and-wait forager when compared the attack aspects of foraging. An unexplored area for future study would be to investigate the differences between sit-and-wait and actively foraging birds in terms of perch location and perch survivorship.

In both South African and North American flycatcher communities the relationships between ecology and morphology are strong and positive and in general, they are concordant. Miles et al. (1987) held that strong relationships between morphology and ecology could be due to relative phylogenetic similarity rather than as a result of adaptive co-evolution. In this study, phylogenetic relatedness within localities is apparent in both behavioral and morphological data. However, the patterns remain when phylogenetic relationships were taken into consideration. With respect to a repeated and hence adaptive aspect to the ecomorphological patterns of these communities, the relationships still may not be as tight as one might expect given the assemblages are composed of groups of sit-and-wait insectivorous passerines. Variation (and statistical distribution) in either behavior, morphology or both due to environmental fluctuations, plasticity or low levels of natural selection on these traits may preclude a tight relationship between ecology and morphology (Weins and Rotenberry 1980). Locality

50 differences may be due to an unmeasured aspect of habitat (prey abundance or distribution) or life history such as migration (Miles, Corbin and Pearson in review).

Further studies including more data on the prey type and abundance or distribution of food items are needed to mete out this confounding factor.

51

CHAPTER 3. CONVERGENCE IN THE ADAPTIVE RADIATIONS OF OLD AND

NEW WORLD FLYCATCHERS

Clay E. Corbin

52 INTRODUCTION

Adaptive radiations are spectacular phenomena in nature. The Australian marsupials, Galapagos finches, New World anoles, Hawaiian honeycreepers, and

Malagasy vangas are all examples of where a single monophyletic clade has evolved into many different unexploited environments. Even more spectacular is when two or more species-rich monophyletic lineages from different continents have independently diversified along common lines of evolution. For example, the sunbirds (Nectariniidae) and hummingbirds (Trochilidae) of Africa and South America, respectively, have coevolved with the nectar-producing plants of those continents. As a result, there is wide variation in the bill morphology among each group that has tracked the evolution of shape in corolla tubes. Another potential example of concordant adaptive radiation is the case of the Old and New World flycatchers. Indeed, the common vernacular names reflect an often-cited example of convergent evolution (Pough et al. 1989) where feeding life-style, wide bills and long rictal bristles are similar features of each lineage.

Surprisingly, what has not been addressed is the extent to which flycatcher lineages exhibit historical concordance in morphological and ecological space.

In both the New and Old Worlds, there have evolved species-rich, independent lineages of flycatchers. I use the term flycatcher to refer to passeriform birds that are primarily insectivorous and comprise the Tyrannidae, Monarchini and Muscicapini.

While the Tyrannid flycatchers share a common ancestor with an Old World group

(Sibley and Alquist, 1990), the radiation has largely occurred since the isolation of South

America (Keast, 1972; Traylor and Fitzpatrick, 1982). Modern day distribution of the family ranges from North to South America. Their ecological counterparts in the Old

53 World, the Muscicapine and Monarchine flycatchers have an Old World origin and coexist across most of the Old World biogeographic realms but are especially abundant in the Afrotropics, Indomalasia and Oceania (see species accounts in Sibley and Monroe

1993).

Similarities between the New and Old World flycatchers in terms of foraging behavior and morphology have been studied to some extent in the past (Keast 1972; Karr and James 1975). However, these analyses have been confined to communities or subsets of the species. While convergence at the community level can be striking (see

Karr and James 1975), the lineages have had different ecological histories (Vaurie 1953;

Keast 1972). For example, the New World flycatchers are thought to have radiated into the under-exploited adaptive zone (sensu Simpson 1953) of sit-and-wait foraging in

South America. In fact, this is described as a key innovation that led to the radiation of the group (Traylor and Fitzpatrick 1982). The Old World lineages coexist with each other and with several other lineages that exhibit this mode of foraging such as bee-eaters

(Meropidae), rollers (Coraciidae), drongos (Dicruridae) and the Australo-Paupan robins

(Petroicidae). Hence, it is possible that the amount of available unexploited habitats or other resources in the Old World were fewer than what was available to the New World flycatchers. In turn, these historical idiosyncrasies may have differentially facilitated or hindered ecological and morphological diversification.

There are multiple resource axes which can be partitioned (e.g. diet, habitat) as assemblage diversity increases. Here the question becomes, is there a specific axis along which the different flycatcher lineages tend to diversify (i.e. MacArthur and Pianka,

1966; Schoener, 1974a; 1974b)? This problem has rarely been addressed incorporating

54 historical information (Losos 1996), and the data that have been published posit different conclusions about the mode and tempo of diversification along resource axes.

Phylloscopus warbler assemblages in the Old World seem to have radiated along a dietary axis early in their evolutionary history and secondarily along a habitat gradient

(Richman and Price 1992; Richman 1996). However, when compared to the New World warbler (Parulidae) radiation, both the tempo and cause of speciation are different. The

New World warblers have diverged more rapidly and possibly due to plumage sexual selection rather than limiting similarities in ecology (Price et al. 2000). Conversely,

Jamaican Anolis lizards have diverged along a habitat axis early in evolutionary history and more recently along a dietary axis (Losos 1992). Furthermore, lineage diversification and age of origin do not necessarily have a linear relationship (Foote 1992; Warheit et al.

1999; Lovette et al. 2002; Corbin and Miles; in review (Chapter 4)).

One way to analyze the comparative ecological diversification of lineages is through ellipse analysis. The technique allows one to assess the qualities of lineage specific trait evolution (Derrickson and Ricklefs 1988; Ricklefs and Nealen 1998).

Briefly, this entails using the parameters of confidence ellipses as metrics of evolutionary diversification. With a representative sample of a species within a clade, one can estimate the tempo (size), direction of diversification (orientation), and the phylogenetic constraint (shape) of their macroevolutionary diversification. This approach is useful when testing hypotheses of evolutionary patterns where there are particular expectations of the attribute space such as character displacement or convergence. Ellipse analysis also has the useful feature that diversification in trait may be used as a standard for comparison to other traits. Hence, the tempo of evolution can be compared between traits

55 (or suites of traits) within or between lineages (Simpson 1944) using this method of analysis.

I use multivariate ordinations and a multivariate extension of bivariate ellipse analysis (Sokol and Rohlf 1995; Ricklefs and Nealen 1998) and measurements of external morphological variation to answer three questions about the evolutionary ecology of independent flycatcher lineages. First, how closely matched are the patterns of morphological covariance among the flycatcher lineages (Corbin Chapter 2; Corbin and Miles Chapter 4; Warheit et al. 1999)? Second, do members of each lineage cluster according to ecomorphological or phylogenetic expectations? In other words, are there

"ecomorphs" within flycatcher lineages (Williams 1983; Leal et al. 2002)? Finally, did diversification occur along unique or common ecological niche axes (MacArthur and

Pianka 1966; Schoener 1974a; Richman and Price 1992; Losos 1996)?

METHODS

Morphology

Species Used in Analyses

For the first question, I used a species list comprised of most of the species of Old

and New World flycatchers in three lineages (Tyrannidae, Muscicapini and Monarchini

(Sibley and Alquist 1990). The world checklist of Monroe and Sibley (1993) is based on

the Sibley and Alquist (1990) tree and was used as a guide for the species pool for

morphological data collection. The Tyrannidae is comprised of four subfamilies, the

Pipromorphinae, Tyranninae, Tityrinae and Cotinginae. For this study, I excluded the

frugivorous Cotinginae.

56 For the second and third questions, I limited my analysis to the species within

Sibley and Alquist’s Tyranninae, Muscicapini and Monarchini. The Tyranninae were then further separated into three subclades: tyranninae, fluvicolinae and elaeniinae

(Traylor and Fitzpatrick 1982; Fitzpatrick 1985). Because Sibley and Alquist's tree agrees with this subclade division, I used genera (and species therein) to test ecomorphological similarity and clustering. See Appendices I and II for species lists.

Morphological Data

My goal when collecting morphological data was to obtain measurements for all species in the three lineages according to Sibley and Monroe (1993). I attempted to obtain measurements for at least 10 individual males for computing a background morphological space (see below) of Tyrant flycatchers (sensu Sibley and Alquist 1990), and at least one individual male specimen for the Pipromorphinae, Tityrinae, Muscicapini and Monarchini. Morphological data for the elaeniine taxa and for the tody-tyrants in the pipromorphinae are from Corbin and Miles (Chapter 4, in review). In some instances, I could not obtain an individual of every species. However, I obtained at least one species from all genera in all lineages except Shiffornis in Tityrinae, Taeneotriccus in

Pipromorphinae; Empidornis, Humboltia and Horizorhinus in Muscicapini; and

Eutrichomyias in Monarchini.

Morphological traits were measured from specimens deposited at the natural history museums of University of Kansas (Lawrence), Louisiana State University (Baton

Rouge), Ohio University (Athens), the Carnegie Museum, the Field Museum (Chicago), the National Museum of Natural History (Washington, D. C.) and the Transvaal Museum

(Pretoria). Measurements were chosen that approximate phenotypic function in the

57 environment (Ricklefs and Travis 1980; Leisler and Winkler 1985; Miles et al. 1987). I used a ruler to measure lengths of the specimen from tip of bill to tip of longest tail feather (total), museum wing chord and the tail. Dial calipers were used to measure the tarsus and toe lengths and four aspects of the bill: its length (tip to naso-frontal hinge), width across the quadrates, width across the nares and bill depth across the naso-frontal hinge. I made two measurements for bill width to capture qualities of gape and tapering

(Hespenheide 1973; Fitzpatrick 1985; Korzoun et al. 2000). All morphological measurements were log10 transformed prior to analysis.

Statistical Analysis

Comparative Morphological Variation.—The radiation of the tyrant flycatchers is

“dominant” in the New World because it has involved the evolution of ecomorphs in the

Americas that are counterparts to several independent lineages on other continents (Keast

1972). In this first part, I wanted to construct a background morphological space

comprised of tyrant flycatchers on which I could superimpose the morphospaces of the

Old World lineages. To compare the morphological variation of the Tyrannidae,

Muscicapini and Monarchini, I first extracted principal components (PCs) from separate

covariance matrices for each group to examine the linear relationships among the

morphological variables for each lineage. Eigenvectors, values and species scores were

estimated using PROC PRINCOMP (SAS 2001). Then, I compared the sign and

magnitude of the coefficients for each lineage along each eigenvector. I also compared

the distributions of scores from the new variables. Relative rates of morphological

evolution in overall body size and then in shape were computed by dividing each

standard deviation of the first two PCs by the lineage's hypothesized relative age of

58 origin. The ages of origin were derived from Sibley and Alquist (1990) DNA hybridization study. I used the delta values (∆T50H) at each corresponding node for each

of the five lineages. The delta values are the differences in temperature from when segments of duplex DNA pairs of the same species and different species denature. As delta value increases, so does the estimated age of origin. Because there is a hypothesized difference in the age of origin within these groups, I wanted to know if the morphological radiation relative to their age of origin is similar among the three groups.

Lineage Differences.—In this aspect of the study, the goal was to compare the morphological divergence among the groups and to identify the traits that were responsible for explaining those differences. I used canonical discriminant functions analysis (CDA) to test for differences in group centroids (PROC CANDISC; SAS 2001).

However, the Tyrannidae is much more species rich than the other lineages (Appendix

II). Many variables (nine), heterogeneous covariance matrices, and the sharply different sample sizes among the three lineages can produce considerable distortions in the Type I error rate (Hakistan et al. 1979) when calculating F statistics. The F statistics in a CDA are more reliable when there is an approximate multivariate normal distribution of observations and homogeneity of variance among group matrices. Hence, for the CDA, I selected genera (and their species) using Sibley and Alquist (1990) as a guide for a representative sampling of each lineage: elaeniinae, fluvicolinae, tyranninae, monarchini and muscicapini. This resulted in a more equitable number of species among the lineages that reasonably approximate the qualities of lineage specific morphological evolution.

The CDA procedure (SAS 2001) finds linear combinations of the original variables that maximally separate the groups. Squared distances (Mahalanobis D2) and F

59 statistics were calculated to test for differences among group means in the multivariate space. I also used resubstitution classification tables from PROC DISCRIM (SAS 2001) to calculate the percent of species that are classified from one group into the other lineages based upon their morphological attributes. By using a prior classification probability that is proportional to group sample size, the resulting misclassification table may help to corroborate the differences observed in the unbalanced CDA.

Ecomorphs.—I determined if the members of each lineage would group together based upon phylogenetic or ecomorphological predictions. Because data on foraging mode is rare for these species in comparison to habitat utilization, I determined if morphology could predict if the species within a genus utilized either Open or Closed habitats. Genera were grouped coarsely into each category according to their member species' habitat designations of Sibley and Monroe (1991). "Open" habitats included scrub, grassland, savannah, forest edge and open woodland designations and "closed" habitats included forest, dense forest, forest undergrowth, cane grass and swamps. I computed geometric grand averages for each genus using the species averages for all morphological data. Then, I subtracted the average from each morphological measurement (Mosimann 1970; Mosimann and James 1979; Klingenberg 1996). Hence,

I consider the patterns in morphological space for this aspect of the study to be relatively free of body size. Using PC-ORD (McCune and Medford 1999) I entered these size-free morphological data into an UPGMA cluster analysis based upon Euclidean morphological distance among the genera. Then I examined, visually, whether genera were grouping according to phylogenetic or ecological expectations. Clusters of genera belonging primarily to the same lineage would be predicted in the first case, and clusters

60 according to either open or closed habitats would verify a prediction of ecology from morphology regardless of phylogenetic affinity.

Axis partitioning.—Because ecological space can be thought of as multidimensional (Hutchinson 1959), a large number of axes would be confusing to interpret accurately. Also, it becomes increasingly difficult to maintain sufficient degrees of freedom with an increasing number of variables (Stevens 1996). Hence, I confine the test of axis partitioning to either a diet (trophic morphology) or habitat (locomotor morphology) axis. The bill morphology (bill length, width 1, width 2, and depth) was used to approximate the dietary differences among the species, while the lengths of the wing, tail, tarsus and toe were used for the habitat preferences (Leisler and Winkler

1985). Again, I computed the geometric average for the morphological values, but this time, for each species in each data set and subtracted the average from the morphological measurement (Mosimann 1970). Hence, I consider the patterns in morphological space for this aspect of the study to be relatively free of body size. I then computed separate

“habitat” and “diet” covariance matrices and their principal components. The first diet and the first habitat PCs were then used to compute a major and minor axis of variation for each lineage (Sokol and Rohlf 1995). Finally, confidence ellipses that include one standard deviation and their parameters of size, shape and orientation were computed for species scores elaeniinae, fluvicolinae, tyranninae, Monarchini and Muscicapini and then for the New and Old World flycatchers combined. The parameters of the confidence ellipses were computed using the program ELLIPSE.SAS of Ricklefs and Nealen (1998, p. 885). Ellipse sizes were compared visually using the standard deviations of the major axis of variation for each group. Ellipse shape and orientations of the morphological

61 disparity were compared in a pair wise fashion among the five lineages with respect to

Old and New World category and then hierarchically between the Old and New World lineages and their corresponding combined values.

RESULTS

Morphological data

I obtained nine measurements on 3256 specimens across 543 Old and New World

species. Sample sizes ranged from one to 41 specimens for each species. Sample sizes,

means and standard deviations for the measurements on the Tyrannidae (sensu Sibley and

Alquist 1990) are presented in Appendix I. Raw values for measurements of the species

in the Monarchini and Muscicapini along with the means for species in the three

tyrannine subclades are presented in Appendix II.

Comparative Morphological Disparity

The first axis in the Tyrannidae is an overall size axis. This is revealed by the roughly

equal magnitudes of positive coefficients in the first eigenvector (Table 3.1). A plot of

the tyrant species along this axis would posit small individuals at one end and larger

individuals at the other (Figure 3.1). The PC explains 80% of the total variation in the

tyrant data (Table 3.2). The second axis explains another 8% of the variation and is a

contrast between bill measurements and the other measures. Hence, individuals with

large bill measurements, mainly bill width 2, and small body measurements would be at

the negative end of the axis whereas species with small, narrow bills and large body

features would occupy the positive end of the axis. The third axis explains 4% of the

variation and describes primarily a contrast between large species with long tails and

62 small leg elements versus small species with short tails and long leg elements. The fourth and fifth axes explain another 3% of the variation. The fourth describes species with relatively long wings and short tails and bills at one end and short wings and long tails and wings at the other. The fifth axis primarily describes differences in bill tapering and depth. Species exhibiting relatively short conical quickly tapering bills (i.e. Anairetes) are posited against those with relatively long, flat bills that taper bluntly near the tip (i.e.

Todirostrum).

As in the tyrannidae, the first axis in the Muscicapini is a size axis. PC1 explains

68% of the variation. PC2 explains 12% of the variation and describes variation between species with wide bills and short tarsi at one end versus species with narrow bills and long tarsi at the other. PC3 explains 9% of the variation and contrasts large species with long wings and tails but short tarsi and flat, narrow bills at one end and at the other, small species with short wings and tails but relatively long tarsi and tall bills. The fourth and fifth axes explain around 6% of the remaining variation. The fourth axis posits species exhibiting short wide bills and long legs and toes against species with long narrow bills and short legs and toes, while the fifth axis is similar to that of the tyrants in that at one end of the axis species exhibit short conical bills and at the other are species with longer bluntly tapered bills.

Like the other groups, a large proportion of the variation in the Monarchini is explained by overall body size. PC1 explains 66% of the variance. PC2 explains 13% of the remaining variance and describes a contrast between species with short tails but long legs and toes at the negative end of the axis and species with long tails but short legs and toes at the positive end of the axis. PC3 explains 10% of the variance and posits species

63 with long tails and narrow bills versus species with short tails and wide bills. PC4 explains 5% of the remaining variation and describes species with deep, narrow bills at the negative end of the axis and tall, narrow bills at the other end of the axis. Finally,

PC5 explains 2% of the variance and describes a contrast between species with a long tarsus and deep, narrow bill but short wing and species with short legs, long wings and a flat wide bill.

The total variance that is explained by the PC model is three times larger in the

Tyrannidae (0.15) than it is for the Muscicapini (0.05) and almost twice that of the

Monarchini (0.08; Table 3.2). Three axes explain 95% of the variance in the tyrants compared to the five axes apiece to explain 95% of the variation in the other groups.

Figure 3.2 depicts the distributions of species scores along the first three PC axes for each group.

With respect to the species along the first PC, the tyrant flycatchers have a much higher standard deviation. However, when the variation is weighted by the sample size, the variation in the tyrants is equal to the muscicapine flycatchers and less than that of the monarch flycatchers (compare standard errors (SE) in Table 3.2). The distributions across the first three PCs and across the lineages are normally distributed except for one case. The species distribution along PC3 in the tyrants is leptokurtotic (g2 = 4.63).

Hence, extreme cases of tail shortening or lengthening are influencing this axis. For

example, Myiornis atricapillus and M. ecaudatus have a small tail to tarsus ratio and are

occupying the extreme positions at the positive end of the axis. On the negative end of the axis are a few birds with a large tail to tarsus ratio: Gubernetes yetapa, Tyrannus

forficatus and T. savanna.

64 A plot of the magnitude and sign of the coefficients from PC1 through PC5 depicts the extent of dissimilarity among the lineages (Figure 3.3). If the groups are tightly convergent with respect to morphological disparity, one would expect that the sign and magnitude of morphological variables to covary between one PC and the next.

Hence, the lines drawn from one PC coefficient to the next within each variable should be relatively parallel and tightly clustered (see the sham variable in Figure 3.3). Because the first PC is an overall size axis in all three lineages, the points for the lineages are all tightly clustered and of the same sign (positive in this case). After the first axis, however, the similarities break down quickly as revealed by the variation in coefficient magnitude and around the origin.

The relative rates of overall body size and shape evolution can be obtained by dividing the standard deviations of the species scores along the first (size) and second

(shape) PCs by the nodal delta value for each group from Sibley and Alquist (1990). For the tyrannidae the rate of overall size evolution is approximately 0.037 (0.35 divided by

9.4) and shape evolutionary rate ~ 0.014 (0.13 / 9.4). For Muscicapini, size rate is ~

0.026 (0.18 / 7.0) and the shape rate ~ 0.011 (0.08 / 7.0). Finally, for the Monarchini, size rate ~ 0.044 (0.22 / 5.0) and shape rate ~ 0.02 (0.1 / 5.0) (Figure 3.4). Hence, the rate of evolution in the monarch flycatchers has exceeded that of the other two groups in terms of overall size and shape. The Tyrannidae have an intermediate rate in both size and shape aspects of evolution and the Muscicapini have, relatively, the slowest rates of the three groups.

65 Morphological Variation in the Five Flycatcher Lineages

The results of the canonical discriminant analysis on the five flycatcher lineages are presented in Table 3.3 and Figures 3.5 & 3.6. The first axis that separates the groups explains 57% of the variation and is an overall size axis. The relatively small value for

Toe reveals its relative unimportance in discriminating among the groups. The second axis explains an additional 31% of the variation and is a contrast between species with a long tarsus and short wings, toes and a narrow bill on the negative end of the axis and species with the opposite features on the positive end. The remaining axes explain the remaining 12% of the variation and are both size related. The group means along all axes are presented in Table 3.4 and the computed distances between the groups (and their F values) are presented in Table 3.5. All groups occupied significantly different aspects of the total morphological space to an alpha value < 0.01. The classification percentages

(Table 3.6) corroborate these differences with low percentages of species being reclassified into another lineage except for the Monarchini. All groups have better than

50% reclassification rates. Given there are five groups, the morphological overlap is relatively low. Interestingly, a scatter plot (Figure 3.6) of the species scores along the first two canonical axes while grouping each lineage into either an Old or New World category, reveals that there may be a lineage dependent size component to the second axis. However, the orientations of the morphological disparity are different with a general positive relationship between Can2 and increasing size in the New World flycatchers (n = 158, b = 0.638, P < 0.001) and a negative relationship in the Old World flycatchers (n = 161, b = -0.581, P < 0.001).

66 Ecomorphs

The results of the cluster analysis reveal that there are elements of both phylogenetic and ecological prediction from morphology (Figure 3.7). The most striking non-phylogenetic grouping based upon similarity is in the middle of the dendrogram within the cluster below the tick mark in Figure 3.7. Several members that utilize closed habitats from Monarchini, Muscicapini and Tyranninae are clustering out as morphological neighbors. Outside this cluster many of the nearest neighbor morphological pairs are mismatched in habitat preference. Some of these cases appear to be phylogenetically generated. For example, Anairetes and Mecocerculus are in the elaeniinae but generally are found in dissimilar habitat types. This could be due to the fact that Mecocerculus may be polyphyletic (Lanyon 1988b) and one or more of the species currently assigned to Mecocerculus are actually phylogenetically more closely related to other members of elaeniine taxa than other members in the genus. Pomerea and Chasiempis (elaeniinae) are likewise phylogenetic neighbor genera but their species are generally found in dissimilar habitats. Other examples may be a reflection of sexually selected characteristics. Gubernetes (fluvicolinae) and Terpsiphone

(monarchini) are not closely related phylogenetically but are morphological nearest neighbors. Most likely, this is due to the long tail feathers exhibited by the species in those genera and this would represent an example of low level of prediction of either phylogeny or ecology from morphology. Hence, at least with respect to closed or open habitat preferences, the clustering in morphological space of flycatcher lineages is largely consistent with the idea of habitat ecomorphs (Williams 1983).

67 Axis Partitioning

The first habitat morphology PC explained 67% of the variation in Old and New

World flycatcher species. The coefficients along the habitat morphology eigenvector are wing = 0.203, tail = 0.72, tarsus = -0.55 and toe = -0.37. Hence the PC axis describes a continuum of species with long wings and tails but short leg elements at the positive end of the axis versus species with the opposite morphological characteristics on the negative end. The first diet morphology axis explains 53% of the variation in Old and New World flycatcher species. The coefficients along the diet morphology eigenvector are bill length

= -0.24, bill width 1 = -0.017, bill width 2 = 0.85 and bill depth = -0.44. Hence, the PC axis primarily describes species with wide, flat bills on the positive end and short, deep bills at the negative end.

The lineage-specific descriptive statistics of the major and minor axes of morphological disparity and the corresponding parameters for confidence ellipses are presented in Table 3.7. T-values between comparisons are presented in Table 3.8. Note that the hierarchical comparisons (i.e. NEW-TYR) are non-independent. Hence, the comparison is meant to be a general description of differences in variation with respect to the lineages. The directional cosine (a12) is one element in the eigenvector of the covariance matrix between the first habitat and diet morphology axes and it estimates the

cosine of the angle between the major axis and the diet morphology axis. The slope

between the variation in diet morphology and habitat morphology is computed by the

-1 equation m = tan(sin [a12]). Graphically, if the ellipse is aligned (orientation

corresponding the group angle in Table 3.7) with the habitat axis, this would support a habitat axis partitioning by the specific lineage as it diversified. On the other hand,

68 partitioning could have taken place along a dietary axis. This would be supported by the alignment of the ellipse along the axis of dietary morphology. Furthermore, if there is a phylogenetic aspect to axis partitioning, the major axis of diversification within a lineage will be similar to the larger clade to which it belongs (Corbin and Miles Chapter 4, in review; Ricklefs and Nealen 1998).

Both Old and New World flycatcher groups seem to have diversified along a habitat axis early in the evolutionary histories of these groups. This is revealed by the low slope values of confidence ellipses (not shown) in Table 3.7. The orientation with respect to the diet morphology axis is slightly higher, but not significant, in the New

World flycatchers (ahd = 0.352) than in the Old World flycatchers (ahd = 0.15) (t = 1.75, P

= 0.08). Within the tyrants there seem to be differences with respect to 1) how the

tyranninae are oriented to the other clades within the New World flycatchers and 2) how

each tyrant clade is oriented with respect to the group as a whole. The tyranninae in both

respects is aligned distinctively along the habitat axis (m = 0.11) whereas the fluvicolinae

and the elaeniinae and then the combined sample are pulled away from the habitat

morphology axis (see respective slopes in Table 3.7 and associated P values in Table

3.8). There are two positive outliers in the tyrannine distribution along the habitat

morphology, the fork-tailed and scissor-tailed flycatchers. If these species are removed

from the ellipse analysis the values of ahd for the tyranninae and the New World flycatchers increase and corresponding apparent differences disappear from Table 3.8.

This also has an effect on the orientation of the entire New World combined sample with respect to the Old World orientation. The t value for differences between group orientation increases and the difference is significant (t = 2.72, P = 0.007).

69 The ellipses are almost twice as long as they are wide in the New World lineages

(shape in Table 3.7). Compare these to the more circular distributions in the Old World lineages: 1.25 (± 0.133) in the Muscicapini and 1.39 (± .163) in the Monarchini. The range of values in the New World lineages is from 1.95 ± (0.305) in the elaeniinae to 3.2

± (0.427) in the tyranninae. Again, upon removal of the two long-tailed Tyrannus spp., the elongate ellipse in the tyranninae drops in its ratio of length to width from 3.2 down to 2.2 (± 0.303), but their removal has little effect on the ellipse shape in the combined

New World sample (compare shapes before and after removal; shapebefore = 1.96 (±

0.156), shapeafter 1.97 (± 0.157)).

To get an idea of the evolutionary rates and changes in tempo of diversification in

these lineages, I divided the standard deviation of the major and minor axes of the ellipse

of each Old and New World lineage and their combined total values by their estimated

relative age of origin. The nodal delta values from Sibley and Alquist (1990) are as

follows: elaeniinae = 5, fluvicolinae = 4.4, tyranninae = 5.7, Muscicapini = 7,

Monarchini = 5, New World lineages combined = 5.7 and most recent common ancestor

to the Old World lineages = 12.8. Hence, with L1 and L2 respectively the resulting

quotients are as follows: ELA = 0.01, 0.006; FLU = 0.02, 0.01; TYR = 0.02, 0.007;

MUS = 0.009, 0.007; MON = 0.019, 0.014; NEW = 0.016, 0.008; OLD = 0.007; 0.005.

The graphical results are shown Figure 3.8. Except for the low values for the elaeniinae,

the rates of evolutionary diversification are relatively low in the ancestral combinations

when compared to their corresponding subclades.

70 DISCUSSION

Ecomorphological concordance is expected when there are similar functional

requirements on the phenotypes of distantly related organisms. Because the species in

this analysis are generally sit-in-wait insectivores, and assuming that morphology

predicts ecology in Old and New World flycatchers (Corbin Chapter 2), one would

predict that there is no difference in the shape or direction of the ecomorphological

diversification if lineages have responded to similar foraging ecological demands (Keast

1972; Ricklefs and Travis 1980; Derrickson and Ricklefs 1989; Ricklefs and Nealen

1998). If there are differences, then the patterns may be explained by different levels of

phylogenetic or ecological constraints on morphological evolution or different levels of

predictive performance of ecology from morphology among the lineages (Weins and

Rotenberry 1980; Weins 1989). The question then becomes: are the historical trajectories

of the lineages concordant with respect to ecology? For example, due to a lower level of competition from distantly related species during the radiation of New World flycatchers, the rates of diversification may be expected to be higher when compared to the Old

World flycatchers.

My first goal of this study was to test whether the morphological variation across independent lineages of flycatchers (Tyrannidae, Muscicapini, and Monarchini) is similar with respect to morphological volume and evolutionary tempo. There is some similarity with respect to overall morphological evolution in these lineages. The main explanatory aspect to both PCA and CDA analyses is overall body size. Divergence in body size has

played a large role in the evolutionary histories of these large clades. To some extent,

this is intuitive because body size is important to so many aspects of a species

71 physiology, ecology and behavior (Schmidt-Nielsen 1984; Calder 1996). In terms of shape, the diversification is largely clade-specific. The functional morphological variables in this study (Leisler and Winkler 1985; Miles and Ricklefs 1994) are likely to have genetic underpinnings (Merila, et al. 1994). These genetic linkages may serve to constrain evolutionary change in any one particular morphological variable (Merila et al.

1994; Diniz-Filho and Sant’Ana 2000). Hence, a size increase or decrease of one morphological trait may cause pleiotropic responses in other morphological traits (Fisher

1930). For example, if diversifying selection of leg length or bill depth impart higher relative fitness to the extreme phenotypes, and those phenotypes are differentially beneficial to populations across a range of habitat types ("open" to "closed"), size changes in other morphological features are likely to ride along with that evolutionary change. Furthermore, it has been shown in studies that concentrate on functionally important traits that evolutionary change is greatest along “lines of least resistance”

(Schluter 1996; 2000). Hence, diversification in shape would have required either a considerable amount of antagonistic natural selection to drive increasing variability across the species in each lineage or lower levels of constraint involved in shape change after species divergence (see Cheverud et al. 1985). This may be the reason that morphological differences generally take the form of variance in overall body size throughout the evolutionary history of a lineage (see Chapter 4)).

When I accounted for size in these lineages' score distributions and their descriptive statistics, the variation in the shape aspects of the morphological space is lineage specific (see Ricklefs and Nealen 1998). In fact, in spite of a predicted size-free ecomorphological convergence, there was little similarity among lineages in the PCA or

72 CDA analyses of species’ morphological distributions. Five main features of the morphological distributions of species revealed in this study lead me to this conclusion.

First, while the first PC is a size axis the variables that are important in explaining the variance along the remaining PCs are dependent upon the lineage. In the first aspect of the study, the eigenvectors after the first PC revealed that the morphological variation is highly lineage specific. Second, in the CDA, the positions of the species within each lineage are clearly different from one another. Third, the second (shape) CD axes in the larger groups comprised of either the Old or New World flycatcher subclades, have entirely different relationships to the first (size) axis. Fourth, the tempo of morphological evolution is not similar among subclades or constant within Old and New World lineages.

Finally, when the morphological disparity of Old and New World lineages was investigated by “constraining” the variance to a couple of axes, significant differences in orientations and shape were revealed. Hence, this points to differential levels of ecological or phylogenetic constraints during the diversification of these lineages.

Theoretical and empirical investigations lead to opposing conclusions as to which resource axis should be partitioned first with an increase in lineage diversity (MacArthur and Wilson 1966; Schoener 1974b; Losos 1992; Richman and Price 1992). The independent early diversifications of these lineages were aligned along a habitat morphology axis as revealed by the alignment of the major axes of diversification for each lineage. And in more recent radiations, this pattern is maintained. However, a plot ellipse shape with respect to the relative age of origin (Figure 3.8) in the subclades reveals that diversification tends to occur in secondary, “special” (Ricklefs and Nealen

1998) components of morphology rather than continuing along a general line of

73 diversification. Hence, it seems that the further along in time from a common ancestor in these lineages, the more likely there is a breakdown of a phylogenetic component to the diversification. Hence, persistence of a clade throughout time may be increasingly less dependent upon the initial phenotypic conditions of the ancestors.

Sit-and-wait foragers theoretically have inexpensive search times when compared to actively foraging birds (Schoener, 1971), and prey size, rather than prey specificity, seems to be a more important aspect to aerial foraging (Hespenheide, 1973; Sherry 1984;

Fitzpatrick 1985). Therefore, if selection is acting in different capacities on biological units (sensu Nemeshkal et al. 1993) in lineages, early morphological radiation in flycatcher lineages may be along habitat axis rather than along a dietary axis. Later local radiations within lineages (Fitzpatrick 1980) may have occurred in response to local prey type and, subsequently, morphological evolution may be responding to changing prey types. On the other hand, if predation in specific flycatcher lineages is related more to prey size rather than taxonomic specificity, partitioning may be along lines of foraging technique (i.e. upward striking, aerial sallying, pouncing etc.). Hence, diversification between diet morphology and habitat morphology covary to a larger extent in some groups (i.e. the tyrants) than in others (i.e. the Old World species). However, there is a general lack of foraging data of these species, especially in the Old World taxa so that attempts to define ecomorphs on the basis of foraging techniques are limited.

Competition can have multiple roles in the diversification of lineages (Lack 1947;

Simpson 1953). Among closely related species or populations, the escape from the negative effects of competition can lead to diversification. On the other hand, a similar or distantly related population or species in a niche can resist the invasion of an inferior

74 competitor. This double faceted characteristic of competition may have played a role in the unique histories of these lineages. Tyrant species are much more numerous than their

Old World counterparts, but on the other hand, the extent to which morphological diversification has occurred since the lineage age of origin is matched or surpassed by the

Monarchine flycatchers. This may be due to the biogeographic qualities of Old and New

World lineage distributions. While there are island representatives of tyrant flycatchers

(i.e. the Caribbean Myiarchus and Tyrannus flycatchers) the Monarchini and to a lesser extent, the Muscicapini are found on islands. Out of 341 species in the Tyranninae

(sensu Monroe and Sibley 1993) 28 or approximately 8% are found primarily on islands whereas ~39% of Muscicapini (46 out of 117 species) are found on islands and almost

80% of the Monarchini (71 out of 89 species) are found primarily on islands. Isolation, historical contingency and competitive ecological situations on islands are thought to foster morphological evolution with respect to their continental counterparts. Examples include the Galapagos finches (Lack 1947; Schluter and Grant 1984), Hawaiian honeycreepers (Lovette, et al. 2001) and Malagasy vangas (Yamagishi et al. 2001). The

Monarchini, while younger than the tyrants, may be another instance where determinants related to island biogeography have facilitated morphological evolution.

There may be more than one way to adapt to an ecological problem. This may be the reason for a general lack of convergence in Caribbean aquatic Anolis lizards (see Leal et al. 2002) and on the scales of magnitude approached in this study. Sit-and-wait insectivory may also be approached by species from different evolutionary backgrounds and that ancestry may constrain species in terms of adaptation to their environment.

Muscicapine flycatchers share ancestry with pouncing bluebirds and ground foraging

75 thrushes, monarchines are close allies of generalist jays and crows, and the tyrant flycatchers share a common ancestor with arboreal sit-and-wait broadbills. Behavioral

(not measured in this study) rather than morphological qualities may facilitate exploitation of aerial foraging prey items. In a comparison between communities of

Southern African and North American flycatchers, the muscicapines utilize pouncing and ground based foraging more than North American flycatchers (Corbin Chapter 2; Corbin unpublished observations on East African flycatchers). This is probably due to the ground foraging and pouncing ancestry of the Old World species.

In conclusion, morphological evolution in independent flycatcher groups is varied and largely lineage-specific. Convergence may be thought of as a “relative” phenomenon

(Keast et al. 1995) in that the magnitude of convergence largely will be dependent upon scale and phylogenetic juxtaposition of the species in the study. The phenomenon is apparent when the similarities of two or more taxonomic units are greater than between their ancestors. In this study, this requirement of convergence is assumed in all analyses but the morphological clustering. Given the vast genetic distance between Old and New

World flycatchers (Sibley and Alquist 1990) and the numerous ecologically different lineages interstitially arranged between the Old and New World Flycatchers, this assumption may be relaxed. Hence, at the lineage-wide level, convergence may be obscured by phylogenetic or behavioral characteristics not measured in this study.

However, the existence of "ecomorphs" based upon a coarse categorization of habitat preference points to convergence in special instances of these species. Future studies using broad scale morphological distributions may need to account for the relative nature

76 of convergence by including another ecologically different taxa such as finch (Schluter

1986) or warbler (Price et al. 2000) clades.

77

CHAPTER 4. MORPHOLOGICAL DISPARITY AND DIVERSIFICATION IN

ELAENIA AND TODY-TYRANT FLYCATCHERS

Clay E. Corbin

and

Donald Miles

78 INTRODUCTION

Phenotypic patterns of macroevolution may be the result of current ecological processes, historical effects, stochastic events or a combination of factors (Ricklefs and

Schluter 1993; Price et al. 2000; Losos 1996; Foote 1995, 1997). It is essential to understand the underlying processes that were important during the radiation of lineages if we are to identify characteristics that are common among evolutionarily successful or persistent lineages (Lovette et al. 2001). The New World flycatchers have been identified as a “dominant family” of suboscine passerines on the South American continent (Keast

1972 and references therein). The ecomorphological space occupied by this group is at least equal the volume occupied by several different lineages in both Africa and Australia

(Keast 1972). The factors responsible for the adaptive radiation of suboscine passerine birds that occurred during the Cenozoic isolation of South America are under considerable debate (see Haffner 1997; Roy et al. 1997; Tuomisto and Ruokolainen

1997). Hence, the goal of this study is not to test for specific mechanisms that led to the suboscine radiations but rather to better illuminate the patterns of flycatcher morphological variability that occurred as a result of that adaptive radiation.

We use an ellipse technique (Derrickson and Ricklefs 1988; Ricklefs and Nealen

1998) in multivariate space (Flury 1988; Klingenberg and Froese 1991) to address questions about the comparative morphological volumes of two species rich flycatcher lineages within the tyrant group; the elaenia and tody-tyrant flycatchers. These lineages are phylogenetically independent from one another, yet have diversified in similar environments and geographic areas. If these groups are convergent we expect their

79 morphological volumes to have similar attributes in terms of size, shape and orientation.

If not, then there could be historical or ecological reasons for the differences. For example, species within the elaenia lineage are known for their ecological generalist foraging behaviors and the tody-tyrants are largely known for their upward striking foraging specialization (Fitzpatrick 1980). Other reasons may be that disparity may correspond positively with relative time since origin or one or more of the component subclades may heavily influence patterns of lineage-wide morphological evolution.

Specifically, the questions we address are: 1) do these species-rich lineages exhibit similar morphological disparity (Gould and Lewontin 1979; Gould 1991; Gould 2002;

Wills et al. 1994; Foote 1997); 2) is the morphological disparity of monophyletic subclades within the lineages predicted by their relative age of origin (see Foote 1996,

1997); and 3) are size, shape and orientation of morphological volume proportionally maintained throughout the history of a lineage (Ricklefs and Nealen 1998)?

The diversification of two or more lineages can be estimated by the differences in their relative morphological disparity. The evolution in these lineages resembles a diffusion of species outward from a centroid in a space described a suite of morphological traits (Derrickson and Ricklefs 1989). Purely random dispersion of a species-rich clade in morphological space would be spherical whereas variation in the rate of evolutionary change in one or more of these traits would alter the sphere into an elongate ellipse (Figure 4.1). Estimation of evolutionary histories of lineages using the relative size, shape and orientation of the ellipses provides metrics for inferences regarding convergent evolution. If we assume that the past evolution in both lineages has occurred as a random process within the morphospace, we predict that the volume

80 occupied by the older tody-tyrant lineage should be greater than the elaenia. Also, a positive correspondence between age of origin and morphological volume is expected to exist when the morphological volumes of tody-tyrant and elaenia lineages are separated into monophyletic subclades. Finally, if there is a lineage-wide phylogenetic constraint, the size, shape and orientations of the hierarchical groupings will be proportional.

METHODS

Morphology

Differences in morphological disparity were quantified using measurements made

on 967 study specimens from 112 species. Skins were obtained from the natural history

museums of the University of Kansas (Lawrence), Louisiana State University (Baton

Rouge) and the Field Museum (Chicago). Total length (distal tip of bill to distal end of

tail) and the lengths of museum wing chord, tail and tarsus were measured with a ruler (±

0.5 mm). Culmen length (from the tip to the nasofrontal hinge), width and depth were

measured with dial calipers (± 0.1 mm). All measurements were log10 transformed prior

to analysis. We attempted to measure at least 10 individuals of each species, however,

sample sizes ranged from 33 for the more common species to one individual for rare

species. Males were primarily used for analysis to avoid problems of sexual dimorphism.

For some species we could not avoid using females. We assume that the morphological variation between sexes is small relative to between species or clades. Morphological

data are in Appendix I. Museum accession numbers can be obtained from the authors.

81 Phylogeny

We used the phylogenetic hypothesis of flycatchers based on morphological characters (Lanyon 1988a, b). For these birds, the tree of Lanyon (1988a, b) differs from the tree derived from DNA hybridization studies of Sibley and Alquist (1990) in that the latter study did not include several genera from the tody-tyrant groups. Hence, we us the inter-lineage relationships hypothesized by the morphological phylogenies to conduct our disparity analyses. However, we used branch lengths from Sibley and Alquist (1990) to estimate age of divergence for the subclades. The best approximate phylogeny (see

Bininda-Emonds, et al. 1999) is presented in Figure 4.2. We assumed the polytomies are soft (Purvis and Garland 1993) and did not attempt to resolve relationships nearer the tips

(McShea 1994). We maintain the polyphyletic nature of the six species of Mecocerculus as in Lanyon (1988a), and we combine Atalotriccus pilaris and the Lophotriccus species

(see Birdsley 1998) (Figure 4.2 and Appendix III).

Subclades

For subclade analyses (see McShea 1994; Wang 2001), tody-tyrants and elaenia were broken up into two and four groups respectively (Figure 4.2). T1 consists of a

“platyrinchus” subclade including three species of Rhynchocyclus, four species of

Tolmomyias, and Onychorhynchus coronatus. T2 includes thirteen species of

Todirostrum, four species of Myiornis, seventeen species of Hemitriccus, two species of

Oncostoma, five species of Lophotriccus, and Poecilotriccus ruficeps. Cnipodectes subbrunneus was not included in either subclade (Lanyon, 1988b), but is included in Ttot

(T). Most of the species in this lineage are considered to have a primarily upward-striking foraging mode, although Hemitriccus has representative perch-gleaning species

82 (Fitzpatrick 1980). The elaenia are broken down hierarchically into three crown groups and a group e4 that is largely comprised of two those crown groups. E1 contains fifteen species of Elaenia, six species of Myiopagis, four species of Tyranniscus, and

Mecocerculus minor and M. calopterus. E2 is defined by the three species of Ornithion, two species of Camptostoma, three species of Mecocerculus (M. poecilocercus, M. stictopterus and M. hellmayri) and Suiriri suiriri. E3 contains Mecocerculus leucophrys, one species each of Capsiempis, Phaeomyias, Nesotriccus and Uromyias (see Roy et al.

1999), and five species each of Serpophaga and Anairetes. E4 is comprised of e2, e3, three species of Pseudocolopteryx and Polystictus pectoralis. Etot (E) is all-inclusive.

Unlike the tody-tyrant lineage above, this lineage exhibits a wide range of foraging

behavior such as perch gleaning in Camptostoma to hover-gleaning and hawking

behaviors found in Phaeomyias and some species of Elaenia (Fitzpatrick 1980).

Overview of Analysis

To assess the extent of morphological disparity within and between the two

flycatcher lineages, we performed separate principal components analyses (PCA) on the

variance-covariance matrices of each lineage. Next we used common principal

components analysis (CPCA) on the total data set with lineage membership as a class

variable. We used a resampling method (Klingenberg 1996, in press; Klingenberg and

Ekau 1996) to compare the angle between lineage PC1 vectors in order to assess whether

there was a common pattern of diversification between the lineages and hence, whether

we were justified in using CPCA. The cosine of this angle is calculated as the sum of the

products of the corresponding PC coefficients (Klingenberg 1996, in press). Then we

used this angle as the test statistic in a bootstrap test to assess whether the lineage

83 diversifications were parallel (the null hypothesis). To construct a null distribution of angles to which we could compare the observed angle, we rotated the coordinates of the morphological data sets to within-group principal components, and hence forcing the PCs for the lineages to be parallel (see Klingenberg 1996). Then we sampled from the “new” data sets 5000 times with replacement and, at each iteration, computed the angle between the lineages. Since the angle between parallel groups should be zero, we located the 95% quantile of the bootstrapped distribution to see if it was smaller than our observed angle

(a rejection of the null). The hypothesis of parallel diversification between these lineages was not rejected (95% quantile = 14.36, observed angle = 8.15, p = 0.854). Therefore, we used CPCA to further characterize the morphological disparities of these lineages.

Common Principal Components

CPCA is a multi-group principal component analysis (PCA) that assumes a common eigenvector among the groups (Klingenberg and Zimmerman 1992). However, it does not assume that each axis explains a similar amount of variance for each group

(Flury 1988; Klingenberg and Zimmerman 1992; Klingenberg 1996, in press). Also,

CPCA maintains the group positions in the space characterized by the morphological measurements. Hence, the variation of one group does not obfuscate the variation in other groups. For comprehensive treatment and reviews of CPCA and multivariate evolutionary allometry, see Flury (1988), Klingenberg (1996), and Arioldi and Flury

1998). We compared the results of the separate PCAs and the CPCA to examine the efficacy of using CPCA in an evolutionary allometry framework (Klingenberg 1996) as well as to better quantify the morphological evolution exhibited within these lineages.

84 Bivariate Ellipses

We constructed bivariate ellipses, including the major and minor axes (Sokal and

Rohlf 1995) for group disparities for all subclades in two analyses, first in size- constrained space (CPC1 and CPC2) and second in size-free space (CPC2 and CPC3).

We computed the Euclidean distance matrices of the species within each group in the two analyses. Median values were then calculated from each group from their respective distance matrices as an index to species packing in morphological space.

We tested the CPC data for normality by comparing the coefficients from the eigenvectors of the CPCA on the log-transformed original data with the means and standard deviations of coefficients from bootstrapped samples (n=1000). There was no difference in the results from the bootstrapped CPCs and the CPCs from the original data; hence, any departures from multivariate normality had little effect on the results of the

CPCA.

We assessed allometry of the original traits in PCs by comparing the observed first coefficients and their corresponding standard errors from 1000 bootstrap iterations to an expected value for isometry (1/√p = 0.4081, where p = number of variables (six in this case)) (Jolicouer 1963; Klingenberg in press). The first PCs describe size variation.

Bootstrapped coefficients in those PCs that are larger or smaller than the expected value have evolved more or less rapidly than body size (Klingenberg in press).

Ellipse analysis

The relationships between two traits or two suites of traits can be characterized by bivariate ellipse parameters (Derrickson and Ricklefs 1988, Ricklefs and Nealen 1998).

85 In particular the size, shape and orientation of an ellipse provide information on variation in morphology during diversification. The differences in these parameters, when the entire lineage is sampled, can then be in interpreted as differences in evolutionary history.

Furthermore, the variation in one variable measures the amount of evolution that has taken place in another variable. We computed 95% confidence ellipses around the bivariate plots of each subclade. First, we used the group scores from CPC1 and CPC2 and second, we used the group scores from CPC2 and CPC3. From the major and minor axes of variation in bivariate space, we computed eigenvalues, and directional cosines from which we could derive the size, shape and orientation of each group’s confidence ellipse. We used the standard deviations of the major and minor axes to evaluate differences in size of the morphological disparity.

The shape of the morphological volume in each group was estimated by the square root of the ratio of the eigenvalues corresponding to the major and minor axes.

One can also estimate standard errors for these values (Sokal and Rohlf 1995; Ricklefs and Nealen 1998), so we used sequential t-tests of pair wise comparisons to determine if orientation and shape were statistically different between subclades. Because we used sequential t-tests, even though the variance parameters (Table 4.3) for the eigenvalues calculated from the major axes were small, we comment on cases where null hypotheses of no difference between groups are rejected at highest alpha values of 0.05 and also

0.01. The alpha value of 0.01 is used as a multiplicity correction of 0.05/5 where 5 is the maximum number of times any one group is included in a pair-wise comparison. We feel that by commenting on significance at both alpha values, we have made an acceptable trade-off between committing type I and type II statistical errors (Bonferroni 1936; Sokal

86 and Rohlf 1995; Perneger 1998). Orientations between groups were also compared using sequential t-tests of subclade directional cosines and their associated standard errors.

Relative Age of Origin and Morphological Disparity

We tested whether the morphological disparity within each subclade was related to its age of origin using non-parametric correlation. A species-rich genus within a subclade (i.e. Hemitriccus) may have an overwhelming influence on the subclade disparity. Furthermore, a lineage may be exceptionally diverse morphologically but have relatively few species, for example, the Malagasy Vangidae (Yamagishi et al. 2001; see also Cibois et al. 2001). In other words, a lineage may be sparsely packed in its morphological space - an expected phenomenon in instances of rapid adaptive radiation or frequent extinction in intermediate morphologies (Findley 1973). In an attempt to partly overcome this potential weakness, we computed the median nearest neighbor distances (NNDs) of each matrix of CPC1 and CPC2 scores for the subclades and lineages. The median NND for each group served as a general index to the amount of morphological evolution that has occurred within the group. Medians, instead of averages were used because they are less influenced by outliers in the NND distributions.

However, the use of average rather than median NNDs for each group did not change the conclusions drawn from the results of a test of the relationship between morphological disparity and age of origin (Corbin unpublished data). The hypothesized ages of origin were taken from Sibley and Alquist (1990) DNA hybridization study. We used the delta values (∆T50H) at each corresponding node in the synthesized phylogeny. The delta

values are the differences in temperature from when segments of duplex DNA pairs of

87 the same species and different species denature. Hence, there is a positive relationship between the delta value and the estimated age of origin.

RESULTS

Patterns of Morphological Variation

The first PC axis explained an equal amount of variance in both the elaenia (75%)

and tody-tyrant (75%) data sets (Table 4.1). The second axis explained 11% and 15% of

the variance in elaenia and tody-tyrant data sets, respectively. Finally, the third axes

explained 8% and 5% of the variance in elaenia and tody-tyrant lineages.

The coefficients of the first eigenvector within each lineage are positive and

roughly equal in magnitude. However, the tail length loads more heavily in the tody-

tyrants than in the elaenia. We interpret this first PC axis in both cases to describe overall

body size. The coefficients for each variable on the second and third axes were strikingly

similar in signs and magnitudes. However, there was a difference in sign between bill

width and depth and increased magnitude in wing length for the elaenia lineage. After

the third eigenvector there has been exactly the same cumulative variance explained in

both lineages. The second axes are shape axes defined mostly by species at one end with

a long tail and tarsus and a small bill, and at the other end, species with a short tail and

tarsus and a large bill. Wing length loaded more heavily in the third elaenia eigenvector

whereas bill depth rather than bill width loaded more heavily within tody-tyrants (Table

4.3).

The CPC coefficients and eigenvalues closely resemble those of the separate

PCAs in magnitude and composition (Table 4.1). This is expected because CPCA finds a

88 common axis of variation and “averages” the differences between the two groups

(Klingenberg in press). Although important information may be contained in the latter vectors of an eigenanalysis (Ricklefs and Miles 1994), we report on only the first three axes from the ordinations. This is because the first three PCs in all ordinations explain around 95% of the variance in each data set. Also in the CPCA, standard errors about bootstrapped (n = 1000) coefficients increase dramatically after the third eigenvector

(Figure 4.3). The first CPC axis describes the common vector of morphological variation in elaenia and tody-tyrant lineages. It explains around 75% of the total morphological variance in both lineages and, as in the separate PCAs, it is a size axis. The second and third axes characterize shape variation in the lineages. The second axis has short tailed and wide billed species at one end contrasting, species with long tails and narrow bills at the other end (Figures 4.4 and 4.5). Along the third axis one finds species with long tarsi and bills at one end and at the other, species with short tarsi and bills (Figures 4.6, and

4.7). The three axes explain 94.15% and 94.39% of the variance in elaenia and tody- tyrant morphological disparities respectively.

The overall morphological disparity is greater in the tody-tyrant lineage

(rangeCPC1 = 0.930, rangeCPC2 = 0.341) than for the elaenia (rangeCPC1 = 0.653, rangeCPC2

= 0.312) (Figure 4.8). Both groups have species that are considered outliers (dark circles in Figure 4.8). The tody-tyrant lineage is skewed (g1 = 0.5615) and leptokurtotic (g2 =

1.274) along CPC1 while elaenia are more evenly distributed along that axis (g1 =

0.3139, g2 = -0.668). Hence, outliers in particular subclades may influence patterns of

lineage-wide morphological disparity. In the separate PCAs, total variance, whether

calculated as the sum of all six eigenvalues (tody-tyrants = 0.0465, elaenia = 0.0366) or

89 the sum of only the first three eigenvalues (tody-tyrants = 0.0441, elaenia = 0.0348) is about 25% larger for the tody-tyrants than in elaenia. Furthermore, if variation due to size (PC1 eigenvalue) is left out of the calculation, there is still about 21% greater variation in shape disparity in the tody-tyrants than what is found in elaenia.

Because there was high similarity in the patterns of covariance of size constrained morphological disparities between elaenia and tody-tyrant lineages, it is not surprising that the results of the analysis on interspecific allometry show similarities as well (Table

4.1). Wing length, tail length and bill width all show positive allometric trends, while tarsus length, bill length and bill depth are negatively allometric. Only the observed coefficient for bill width in elaenia is not outside the standard error of the expected value.

Ellipse Analysis

From the species scores along the first three common principal components we plotted against each other, 1) the first and second (size-constrained) and, 2) the second and third (size free) CPC scores for all species (see Jolicoeur 1963; Burnaby 1966;

Klingenberg 1996). These plots were then used in the ellipse analysis. The entire results of the ellipse analysis are presented in Appendix IV. Of primary importance here are the size, shape and orientation statistics of each subclade.

Ellipse Size

If morphological radiation occurs as a function of the time since origin, we predict a smaller morphological disparity among the younger lineages. The standard deviations of the major and minor axes estimate the size of a group’s morphological volume. In the elaenia lineage, with exception of e2, this pattern holds up in size-constrained space.

90 However, in the size-free space defined by the second two CPCs, there is considerable departure from what is expected from a random model of morphological evolution. Based on Sibley and Alquist (1990) branch lengths, an expected ranking in standard deviation from largest to smallest were E, e1, e4, while e2 and e3 should be similar. The observed ranking in size free space is e1, e2, e4, E, e3. Similar patterns can be noticed in the tody- tyrants. While the younger groups follow an expected trend in size-constrained space, in size-free space, t2 is less disparate than what one would expect from a linear model of morphological evolution. Further patterns of morphological evolution within these lineages can be described using bivariate scatter plots with CPC1 and CPC2 as the x and y variables respectively and confidence ellipses surrounding the group scores along those axes. The lineage scatter plots are displayed in Figures 4.3-4.6 and corresponding t- values for subclade pair wise comparisons in shape and orientation are listed in Table 4.2.

For ease of visual comparison, species scores in Figures 4.3-4.6 are plotted with axes of the same dimensions.

Ellipse Shape

There are no differences in the ellipse shape statistics in either CPC1 vs. CPC2

(size-constrained) or CPC2 vs. CPC3 (size-free) space of elaenia and tody-tyrant lineages as a whole (t = 0.766, p = 0.445, t = 558, p = 0.578, respectively). Many of the ellipse shape characteristics are maintained throughout the elaenia lineage. In the size- constrained space, only one pair wise comparison (e1-e4) is significant at the alpha level of 0.01. The volume of group e1 is more elongate than that of e4 (t = 2.71, p = 0.009).

Also, one (E-e1) is significant at the alpha level of 0.05 (t = 2.37 p < 0.02). Again, e1 has the more elongate disparity. So in the size-constrained space, except for the group that

91 largely consists of species in the genus Elaenia (e1), all ellipses are essentially proportional. Only one ellipse shape comparison (e2-e4, t = 2.15, p = 0.038) out of the 14 is significant in the size-free space. The group e2 is more elongate with respect to axis 3 than is e4. The size-free comparison between e1 and e4 approaches significance but only with respect to an alpha value of 0.05 (t = 1.98, p = 0.0523), the ellipse of e1 being more elongate than that of e4.

Ellipse shape differences were found in both size-constrained and size-free morphological space of the tody-tyrants. There was a significant difference between the total lineage (T) and the group that is comprised largely of Todirostrum and Hemitriccus species (t2). Also this last group has a different shape ratio closer to unity in size-free space in comparison to the other subclade in the linage (t = 2.05, p = 0.0449).

Ellipse Orientation

The orientation statistics reveal fewer similarities among the subclades. In the size-constrained morphological space of the elaenia group, three of the 11 pair wise comparisons are significant to a maximum alpha value of 0.01. The elaenia as a whole differs from e4 in that e4 is more negatively oriented (t = 2.94, p = 0.004). The group that contains the genus Elaenia (e1) has a slightly positive aspect to its orientation while the sparsely packed group e2 is negatively oriented (t = 2.897, p = 0.007). At the alpha value of 0.05, e2 is negatively oriented with respect to the total lineage orientation (t = 2.46, p =

0.017). Interestingly, in the size free space, there is a significant difference in the orientations of elaenia and tody-tyrant lineages. While both are positive, the elaenia is more aligned with CPC3 than the tody-tyrants (t = 3.517, p < 0.001), clade e1 (t = 2.87, p

= 0.005) and e2’s more inclusive group e4 (t = 1.986, p = 0.0502). We state the

92 significance of the latter comparison with caution due to a rounding down of the p-value and the problem of multiplicity. Finally, group e1 is oriented slightly less positively than e3 in the size free morphological space (t = 2.05, p = 0.047).

Age of Origin and Morphological Disparity

The median nearest neighbor distances were plotted against melting temperatures from a DNA-DNA hybridization study (Sibley and Alquist 1990) to test for a positive relationship between the age of origin and species packing. Delta values and median

NNDs for both size constrained (CPC1 and CPC2) and size free (CPC2 and CPC3) morphological spaces are presented in Table 4.3. We found no significant correlation between age of origin and disparity in size constrained (Spearman’s rho = 0.120, p =

.606) or size free (rho = 0.217, p = .606) morphological spaces.

DISCUSSION

Use of CPCA and Separate PCAs

The separate PCAs show that in each group, PC1 accounts for approximately 75%

of the total variance. This similarity in the amount of explained variance extends also to

PC2 and PC3. In these situations, CPCA is preferred to separate PCAs because standard

errors that are computed from the extracted components are smaller (Arioldi and Flury

1998). So, in data sets with large variability, a CPCA may be preferred. Furthermore, the

CPCA extracts unrelated components from the original data with the added benefit of

maintaining group positions in multivariate space. Here, it would have been difficult to

visualize the lineage volumes with respect to one another if we did not use the CPCA

model. In addition, the technique lends itself nicely to accounting for body size in

multiple group comparisons (Klingenberg in press; this study). We encourage the use of

93 CPCA with a resampling component over separate PCs in situations of comparative analysis of highly variable, multi-group, morphological space.

Patterns of Morphological Evolution

The lineage-wide variation for both elaenia and tody-tyrants parallel an overall body-size axis (Figures 4.4 and 4.5). This suggests the early morphological radiation in these lineages occurred along a size axis. Tody-tyrants are more diverse in size than elaenia, which may suggest a constraint in elaenia morphological evolution. However, when one examines the contributions of each subclade to the variance in size, two patterns emerge. First, the greater variation in the tody-tyrants results from a few divergent taxa (i.e. Myiornis or Rhynchocyclus (Figure 4.4)) in the subclades. Second, the size variation in elaenia is evenly distributed across the subclades.

The size component to these data follows common scaling phenomenon observed in passerine birds (Blackburn and Gaston 1994a; 1994b). Illuminating the mechanisms responsible (deterministic or stochastic) for the phenomenon is a major problem of study in evolutionary ecology (Brown and Maurer 1986, 1987; Maruer 1998a, b). An analysis of each mechanism is beyond the scope of this study. However, it is important to comment on the overwhelming explanatory power of size in these data. Because we used functional morphological variables in this study (Leisler and Winkler 1985, Miles and

Ricklefs 1994) these covariances are likely to have genetic underpinnings (Merila et al.

1994). This size-related genetic correlation along with other factors (i.e. structural, ontogenetic, physiological, etc.) may serve to constrain evolutionary change in any one particular morphological variable (Merila et al. 1994; Diniz-Filho and Sant’Ana 2000). A

94 selection for size increase or decrease of one morphological trait may cause pleiotropic responses in other morphological traits (Fisher 1930). Furthermore, it has been shown in studies that concentrate on functionally important traits that evolutionary change is greatest along “lines of least resistance” (Schluter 1996). Hence, differences in shape traits such as what is represented in CPC2 and CPC3 would have required either 1) a considerable amount of antagonistic natural selection to drive increasing variability across the species in each lineage, or 2) a relaxation of the constraints involved in evolutionary shape change after species divergence (see Cheverud et al. 1985).

Trait allometry with respect to overall size was apparent when subclade information was ignored (PC1s in Table 4.1). The lineage wide patterns of short tarsi and wide bills correspond to the predictions of functional morphology in flycatchers

(Fitzpatrick 1985; Erard 1987; Korzun et al. 2000). The patterns of bill allometry reflect the generalist foraging behavior for many of the species within the lineage (Williamson

1975; Fitzpatrick 1980; Marini and Cavalcanti 1998). When subclade information is taken into account in the ellipse analysis, much of the lineage-wide variation was explained by the unique evolutionary trends observed in the less inclusive groups. For example, the subclades t1 and t2 occupy different ends of the size and shape components

(discriminant functions analysis (DFA) on CPC1-3: Wilk’s Λ = 0.225, p < 0.0001). The subclade t2 exhibits similar characteristics in disparity as the tody-tyrant lineage as a whole, while t1 differs in orientation. This is also shown in the elaenia with the positions of e1 and e2 (DFA, CPC1-3: Wilk’s Λ = 0.55, p = 0.0002). So, while size variation is apparent in these lineages with respect to particular traits, the pattern can be explained by the relative size, position and orientation of ellipses for each subclade within the lineages.

95 In other words, the factors that seem to have been important during the evolution of one clade are not necessarily the same factors important to another. Significant differences in the shape and orientation of the subclade disparities lend evidence to this conclusion.

Size differences in morphological volumes of Tody-tyrant and Elaenia flycatcher

lineages.

Relative Age of Origin

We found no evidence for a positive relationship between disparity and age of origin. According to recent DNA hybridization data and a South American – African tectonic plate calibration (Sibley and Alquist 1990) the subfamily pipromorphinae, which includes members of the tody-tyrant lineage in this study, may be up to 3.7 million years older than the elaenia taxa. So, one could argue that the morphological disparity of the tody-tyrant lineage is larger than that of the elaenia simply because the lineage is older.

However, the lack of a positive relationship between morphological disparity and the age of origin reveals that the tempo of morphological evolution was not constant when subclade information was taken into account.

Current Ecological or Historical Factors

Ecological mechanisms including limiting similarity and can promote the increase of morphological variability in particular lineages. These include taxon-cycling (see

Wilson 1961; Ricklefs and Cox 1972; and Jones et al. 2001) and community-wide character displacement (see Lack 1947; Strong et al. 1979; Ben-moshe et al. 2001).

Taxon-cycling has been proposed as a mechanism leading to the diversification of species belonging to the tody-tyrant group (Fitzpatrick 1976). The isolation, evolution and competitive exclusion of similar species may have been facilitated by cyclic patterns of

96 wet and dry habitat fragmentation during South America’s geological past. Also, geographical separation by the complex Amazonian river systems of South America coupled to a competitive resistance of invasion has also been proposed as an important factor in the maintenance of species segregation within Hemitriccus (Cohn-Haft 2000).

However, this segregation may be related to genetic and song variation rather than to morphological variability (Cohn-Haft 2000). Perhaps the most inclusive ecological explanation of morphological variability of these lineages to date is the hypothesis of past ecological opportunities (Keast 1972). A lack of insectivorous competitors during the radiation of these birds in South America may have led to competitive release of species from ancestral suboscines in South America. However, much of the specific ecological nature of morphological evolution is beyond the scope of this study and is currently being considered by other researchers (Johnson 2001, 2002).

The patterns of morphological disparity observed in these flycatcher lineages show three trends. First, the tody-tyrant lineage as defined by Lanyon (1988b) occupies a larger morphological volume than the elaenia lineage. However, the positions in the total morphological space of elaenia may be influenced by particular subclades. For example group e2 (the smallest subclade in the elaenia comprised of Ornithion, Camptosoma, three species of Mecocerculus and the Suiriri species) had a larger morphological disparity than what would be predicted by its age of origin or number of species. The tody-tyrant lineage is comprised largely of two groups – an under-dispersed clade

(Todirostrum, Poecilotriccus, Myiornis, Hemitriccus, Oncostoma and Lophotriccus) and the other hyper-dispersed group (Rhynchocyclus, Tolmomyias, Onychorhynchus and

Platyrhynchus). Hence, second, we conclude that there is no relationship between the

97 morphological disparity in these groups and their estimated age of origin. This conclusion is dependent upon the phylogenetic hypothesis of Lanyon (1988a, b) and the concomitant relative ages of origin estimated by Sibley and Alquist (1990). However, Foote (1992,

1996) has shown that this relationship between morphological disparity and age of origin does not necessarily increase in a linear fashion over large time scales. His conclusions are based in part upon the evidence of different rates of macroevolutionary response to ecological opportunities. Third, we find that at least in thee two lineages of flycatchers the factors important to the morphological disparity of clades of organisms are not common. If ecological factors such as competition are important in the morphological disparities of these lineages, we expected to observe the maintenance of shape proportions and parallel orientations among the subclades. However, while we did observe a general proportionality of shape in the ellipse statistics across subclades, the patterns of ellipse orientation were not maintained. Hence, the decoupling of these ellipse characteristics lead us to conclude that stochastic events (i.e. geographic vicariance, weather) or historical factors (i.e. extinction, speciation and species dispersal) are at least equally if not more important as current ecological factors that are suggested (Fitzpatrick

1976) leading to the morphological patterns that we observe in these flycatcher lineages.

98

CHAPTER 5. GENERAL CONCLUSIONS

Clay E. Corbin

99

In this study I set out to compare and contrast the patterns of macroevolution within and among flycatcher clades of the world. To do this, I characterized species’ relative positions in space using measurements on the morphology and foraging behavior for 632 species of Old and New World flycatchers. There are four general conclusions that I can draw from the tests of hypotheses above.

1. As expected from an adaptive form-function hypothesis, there is a significant positive relationship between the morphology and foraging behavior in Old and New World flycatchers. Furthermore, these relationships are concordant between North American and African flycatcher communities. This is the first test of ecomorphological concordance between flycatcher communities while taking into account the phylogenetic relationships among the community members.

2. Convergence more likely should be attributed to special cases between ecomorphs within Old and New World flycatchers and not necessarily be attributed to the clades as a whole. These findings may reflect the general observation that Tyrannidae is a dominant family in South America in that the single clade occupies a manifold of ecological opportunities that are counterparts to several independent clades of the Old World (Keast

1972). However, this study quantifies the ecological “dominance” of the Tyrannidae.

3. The relationship between morphological disparity and the age of origin (if any) is not linear. This can be explained by particular over or underdispersed subclades that offset the relationship. The non-linear relationship can be seen when comparing Old vs. New

100 World flycatchers (Chapter 3) as well as within the New World clades of elaenia and tody-tyrants.

4. The major axis of diversification is not always maintained by younger subclades.

Hence, ecological interactions can outweigh historical factors in explaining the patterns of the morphological evolution of flycatchers. There are three lines of evidence that support this conclusion. First, there is an adaptive nature of ecomorphological relationships between the North American and Southern African flycatcher communities.

Second, at a course habitat level, genera group toward habitat preference and away from phylogenetic affinity. Finally, as above, the major axis of diversification in these clades is not necessarily the same axis of diversification of the hierarchical grouping above it.

Hence, this study is important in that it reveals the bipartite nature of ecomorphological evolution in large, extant (cf. Foote 1997) clades of passeriform birds.

Furthermore, this work is important in shedding light on several aspects of evolutionary ecology. By providing a clearer view of the purported convergence between Old and

New World flycatchers (Pough 1985)., testing (and confirming) ecomorphological concordance while taking into account phylogenetic affinity (Miles et al. 1987) and testing the hypothesis that morphological variation should increase with increasing age of origin (Foote 1997), I hope this work will provide insight into issues fundamental to both ecological and evolutionary biology.

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Table 2.1. The cross tabulated data entered into the correspondence analysis on search behavior. The cells are the enumerated percentages of each species with respect to where the bird was searching for food. C = under the canopy, O = in the open. PH1 = Lowest perch height. PH5 = Highest perch height. See text for category values.

Species CPH1 CPH2 CPH3 CPH4 CPH5 OPH1 OPH2OPH3 OPH4 OPH5 Contopus pertinax 0.00 0.00 0.00 0.00 0.03 0.00 0.00 0.12 0.03 0.82 Contopus sordidulus 0.04 0.10 0.07 0.04 0.16 0.01 0.02 0.01 0.07 0.48 Contopus virens 0.01 0.03 0.03 0.02 0.67 0.03 0.00 0.02 0.00 0.19 Empidonax fulvifrons 0.05 0.10 0.10 0.14 0.31 0.01 0.05 0.02 0.01 0.21 Empidonax traillii 0.09 0.15 0.06 0.02 0.01 0.28 0.13 0.04 0.05 0.17 Empidonax virescens 0.04 0.13 0.16 0.06 0.54 0.00 0.00 0.01 0.00 0.06 Myiarchus cinerascens 0.26 0.10 0.11 0.02 0.03 0.19 0.16 0.09 0.01 0.03 Myiarchus tuberculifer 0.12 0.03 0.17 0.10 0.37 0.00 0.00 0.02 0.01 0.18 Myioborus pictus 0.13 0.05 0.17 0.31 0.32 0.02 0.00 0.00 0.00 0.00 Myiodynastes luteiventris 0.13 0.07 0.09 0.07 0.53 0.00 0.00 0.00 0.00 0.11 Pyrocephalus rubinus 0.09 0.23 0.06 0.03 0.02 0.07 0.09 0.03 0.06 0.32 Sayornis nigricans 0.29 0.21 0.04 0.02 0.02 0.31 0.02 0.00 0.01 0.08 Sayornis phoebe 0.07 0.13 0.03 0.00 0.04 0.09 0.17 0.13 0.21 0.13 Tyrannus tyrannus 0.01 0.02 0.00 0.02 0.01 0.09 0.00 0.29 0.05 0.51 Tyrannus verticalis 0.00 0.01 0.00 0.00 0.00 0.09 0.33 0.14 0.00 0.43 Tyrannus vociferans 0.00 0.03 0.00 0.00 0.05 0.22 0.15 0.14 0.03 0.38

121 122 Table 2.2. The coefficients along the eigenvectors, the eigenvalues and the cumulative explained variance for the correspondence analysis on the enumerated, cross-categorized variables of perch height and foraging location in New World flycatchers. C = under canopy, O = in the open. The perch height variables run from low (1) to high (5).

Variable Search1 Search2 Search3 UPH1 0.20 -0.84 0.32 UPH2 0.01 -0.54 0.08 UPH3 0.67 -0.23 0.14 UPH4 0.96 -0.10 0.54 UPH5 1.03 0.32 -0.31 OPH1 -0.58 -0.83 0.03 OPH2 -0.80 -0.36 -0.74 OPH3 -0.87 0.39 -0.07 OPH4 -0.66 -0.16 -0.33 OPH5 -0.50 0.61 0.21 Eigenvalue 0.49 0.28 0.10 % Cum. Var. 43 68 76

123

Table 2.3. The foraging percentages for each foraging mode used in the correspondence analysis of attack behavior for Old and New World Flycatchers. Abb. = abbreviations used in figures. P = pounce, S = standpick, H = hover-glean, F = flycatch. Sample sizes (n) are the sequential sample sizes of combined totals of personal and Fraser’s (1983) observations. * = Old World species.

Species Abb. n P S H F Batis species* BAT 195 0 11 61 28 Bradornis mariquensis* BM 77 79 21 0 0 Melaenornis pammelina* MP 270 69 2 2 27 Muscicapa striata* MS 81 15 8 2 75 Myioparus plumbeus* MPb 51 0 26 70 4 Sigelus silens* SS 67 72 3 0 25 Terpsiphone viridis* TV 71 0 16 52 32 Contopus pertinax cp 37 0 0 0 100 Contopus sordidulus cs 205 0 1 7 92 Contopus virens cv 117 0 0 19 81 Empidonax fulvifrons ef 61 0 0 42 58 Empidonax traillii et 137 4 0 38 58 Empidonax virescens ev 90 0 8 34 58 Myiarchus cinerascens mc 69 0 2 16 82 Myiarchus tuberculifer mt 128 0 3 55 42 Myioborus pictus mp 11 0 29 11 60 Myiodynastes luteiventris ml 28 7 0 11 82 Pyrocephalus rubinus pr 106 4 1 10 85 Sayornis nigricans sn 143 6 5 9 80 Sayornis phoebe sp 86 34 3 6 57 Tyrannus tyrannus tt 110 1 3 1 95 Tyrannus verticalis tve 205 4 0 2 94 Tyrannus vociferans tvo 109 12 1 4 83

124 Table 2.4. The coefficients along the eigenvectors, the eigenvalues and the cumulative explained variance for correspondence analysis attack mode.

Variable Attack1 Attack2 Attack3 Pounce 2.48 0.31 -0.52 Standpick 0.13 1.62 3.53 Hoverglean -0.81 1.58 -0.98 Flycatch -0.30 -0.74 0.07 Eigenvalue 0.54 0.33 0.08 % Cum. Var. 57 92 100

125 Table 2.5. Means of morphological measurements for flycatcher species used in this study. L = locality, n = sample size, Tot = total, Tar = Tarsus, BW1 = bill width across the quadrates, BW2 = bill width across the nares, BD = bill depth.

Species L n Tot Wing Tail Tar Toe BL BW1 BW2 BD Contopus pertinax AZ 11 171 101.6 86.2 16 15 22 11.8 8 6.1 C. sordidulus AZ 20 144 85.6 69.7 13 13 18 9.6 6.8 4.6 Empidonax fulvifrons AZ 20 109 58.2 51.3 14 11 13 7 4.6 3.3 Myiarchus cinerascens AZ 20 192 98 97.8 22 17 24 11.7 7.3 6.3 M. tuberculifer AZ 20 159 77.7 76 19 14 21 10.3 7 4.9 Myioborus pictus AZ 1 116 64 59 16 13 12 6.6 4.3 2.7 Myiodynastes luteiventris AZ 20 190 113.2 88.2 19 20 26 16.6 10.2 8.3 Pyrocephalus rubinus AZ 20 132 80.8 62.6 16 14 17 8.7 5.5 4.1 Sayornis nigricans AZ 20 165 89.3 81.7 18 15 19 8.9 5.7 4.4 Tyrannus verticalis AZ 20 205 128.5 99.4 18 20 24 14.8 8.6 7 T. vociferans AZ 20 209 128.4 96.9 19 20 24 15.5 9.2 7.1 Contopus virens OH 20 145 82.5 69.3 13 13 18 10 6.5 4.6 Empidonax traillii OH 29 133 69.2 60 16 14 16 8.8 5.9 4.3 E. virescens OH 20 134 72 62.2 15 13 17 9.3 6.4 4.2 Sayornis phoebe OH 20 161 84.9 74.9 17 15 19 9.2 6 4.5 Tyrannus tyrannus OH 20 197 117 87.8 19 19 21 13.5 8.4 6.7 Batis species SA 3 123 58.4 46.2 18 8.9 14 9.2 4.5 7.4 Bradornis mariquensis SA 2 159 81.5 79 22 17 17 9.6 4.6 4.3 Melaenornis pammelina SA 20 195 105.4 98.4 22 20 19 10.9 5 5 Muscicapa striata SA 30 144 87.4 66 14 11 17 8.5 4.7 4.7 Myioparus plumbeus SA 1 129 73 66 18 14 15 8.4 4.4 3.7 Sigelus silens SA 1 177 87 82 24 20 20 10.5 4.8 4.7 Terpsiphone viridis SA 1 191 89 115 15 14 19 9 6 4.4

126 Table 2.6. The coefficients, eigenvalues and percent cumulative variance explained for the principal components analysis (PCA) on the covariance of log10 transformed morphological variables in Old and New World flycatcher species. Bill Width 1 = bill width across the quadrates. Bill Width 2 = bill width across the nares.

Variable PC1 PC2 PC3 Total 0.31 -0.233 0.198 Wing 0.352 -0.099 0.506 Tail 0.324 -0.328 0.243 Tars 0.202 -0.497 -0.427 Toe 0.312 -0.34 -0.485 Bill Length 0.304 0.064 0.032 Bill Width 1 0.399 0.234 -0.112 Bill Width 2 0.351 0.588 -0.399 Bill Depth 0.403 0.248 0.234 Eigenvalue 0.061 0.007 0.003 % Cum. Var. 79 89 93

Table 2.7. Summary of the canonical correlation analysis (CCA) on search behavior.

Canonical Canonical Canonical Variate Correlation R2 F df*P** Search Behavior and Morphology Original Data 1 0.81 0.66 2.6 9 0.03 2 0.62 0.38 1.7 4 0.18 3 0.24 0.06 0.72 1 0.41

Search Behavior and Morphology Contrast Data 1 0.79 0.63 1.18 9 0.37 2 0.52 0.28 0.64 4 0.64 3 0.11 0.01 0.09 1 0.77 *df = (p + 1 - i)(q + 1 - i), df = degrees of freedom, p = no. morphological variables, q = no. behavioral variables, i = ith variate. ** Tests the null hypothesis that the row and all that follow are zero

127 Table 2.8. Summary of the canonical structure of morphology and search behavior in New World flycatchers.

Correlations between the Correlations between the search variables morphological traits and their and their canonical variables canonical variables Search CA Canonical Axis 1 Morphological Canonical Axis 1 Axis (search) PCA Axis (morphology) 1 0.14 1 -0.4 2 -0.94 2 0.92 3 0.3 3 0.32

Table 2.9. Summary of the canonical correlation analysis (CCA) on morpohology and attack behavior in Old and New World flycatchers.

Canonical Canonical Canonical Variate Correlation R2 F df*P** Attack Behavior and Morphology Original Data 1 0.83 0.7 4.3 9 0.0005 2 0.57 0.33 2.01 4 0.11 3 0.1 0.01 0.18 1 0.68 Attack Behavior and Morphology Contrast Data 1 0.89 0.78 3.6 9 0.003 2 0.4 0.16 0.65 4 0.63 3 0.04 0.002 0.03 1 0.86 *df = (p + 1 - i)(q + 1 - i), df = degrees of freedom, p = no. morphological variables, q = no. behavioral variables, i = ith variate ** Tests the null hypothesis that the row and all that follow are zero

128 Table 2.10. Summary of the canonical structure of morphology and attack behavior in Old and New World flycatchers.

Correlations between the attack Correlations between the variables and their canonical morphological traits and their variables canonical variables Canonical Axis Attack CA Canonical Axis Morphological 1 Axis 1 (attack) PCA Axis (morphology) Attack Original Data 1 0.74 1 -0.23 2 -0.61 2 0.99 3 -0.25 3 0.28 Attack Contrast Data 1 0.8 1 0.68 2 0.84 2 -0.15 3 -0.48 3 -0.86

Table 2.11. Results of the ANCOVA testing for differences in Locality means in the foraging variables with morphological covariates.

Source SS I F P SS III F P R2 Dependent Variable = Attack1 (CA axis1) Locality 2.20 7.70 0.01 0.34 1.19 0.29 0.56 Morph1 1.08 3.80 0.07 1.16 4.04 0.06 Morph2 3.16 11.02 0.004 1.70 5.91 0.03 Morph3 0.23 0.81 0.38 0.23 0.81 0.38 Dependent Variable = Attack2 (CA axis2) Locality 3.67 27.39<0.0001 2.24 16.70 0.0007 0.70 Morph1 1.29 9.63 0.006 0.88 6.60 0.02 Morph2 0.07 0.50 0.49 0.03 0.23 0.64 Morph3 0.64 4.80 0.042 0.64 4.80 0.04

129

Table 2.12. Attack rates and standard errors (in parentheses) based upon grand averages of species with n individuals drawn from the total data set. Each individual is comprised of three sequential search times. Attack rate is the mean number of times per second an individual attempted a prey capture.

Species n(3) Attack Rate Contopus pertinax 6 0.029 (0.004) Contopus sordidulus 26 0.050 (0.007) Contopus virens 16 0.067 (0.021) Empidonax fulvifrons 14 0.126 (0.043) Empidonax traillii 35 0.063 (0.005) Empidonax viresens 11 0.068 (0.008) Myiarchus cinerascens 16 0.060 (0.008) Myiarchus tuberculifer 23 0.088 (0.011) Myioborus pictus 24 0.451 (0.034) Myiodynastes luteiventris 6 0.027 (0.006) Pyrocephalus rubinus 20 0.044 (0.007) Sayornis nigricans 22 0.063 (0.013) Sayornis phoebe 14 0.049 (0.009) Tyrannus tyrannus 12 0.039 (0.008) Tyrannus verticalis 14 0.016 (0.003) Tyrannus vociferans 9 0.016 (0.003) Bradornis mariquensis 5 0.050 (0.010) Melaenornis pammelina 4 0.032 (0.004) Sigelus silens 2 0.026 (0.011) Batis molitor 2 0.123 (0.034)

130 Table 3.1. Coefficients of the principal components analysis on separate covariance matrices.

Lineage Variable PC1 PC2 PC3 PC4 PC5 Total 0.336 0.214 -0.244 0.035 -0.029 Wing 0.362 0.200 -0.015 -0.864 0.032 Tail 0.374 0.366 -0.658 0.292 0.087 Tarsus 0.171 0.467 0.509 0.180 0.157 Toe 0.245 0.216 0.372 -0.007 -0.066 Bill Length 0.356 -0.034 0.291 0.284 0.424 Tyrannidae Bill Width 1 0.349 -0.227 0.135 -0.009 -0.497 Bill Width 2 0.387 -0.651 -0.049 -0.043 0.463 Bill Depth 0.356 -0.198 0.067 0.229 -0.566 Total 0.359 -0.039 -0.260 0.043 0.048 Wing 0.319 0.012 -0.356 0.107 -0.346 Tail 0.434 -0.128 -0.589 -0.146 0.257 Tarsus 0.301 -0.592 0.387 0.333 -0.054 Toe 0.272 -0.261 0.231 0.310 0.094 Bill Length 0.292 -0.005 0.221 -0.334 0.642 Muscicapini Muscicapini Bill Width 1 0.372 0.390 0.058 0.314 -0.334 Bill Width 2 0.230 0.639 0.254 0.283 0.302 Bill Depth 0.373 0.065 0.371 -0.684 -0.431 Total 0.374 -0.257 -0.250 0.012 -0.003 Wing 0.371 0.069 -0.094 0.104 -0.433 Tail 0.402 -0.466 -0.552 -0.020 0.224 Tarsus 0.220 0.601 -0.192 0.394 0.436 Toe 0.276 0.402 -0.136 0.118 -0.298 Bill Length 0.366 0.171 0.223 0.034 -0.082 Monarchini Bill Width 1 0.283 -0.058 0.263 -0.138 -0.492 Bill Width 2 0.237 -0.369 0.579 0.599 0.222 Bill Depth 0.407 0.136 0.337 -0.664 0.430

131

Table 3.2. Descriptive statistics of the principal components analyses on separate covariance matrices.

Lineage Descriptive Statistic Cum. Total PC SD SE Kurt Skew Eigen Var. Var. 1 0.35 0.020 -0.67 0.31 0.120 0.80 0.15 2 0.13 0.007 0.46 -0.19 0.016 0.91 3 0.08 0.005 4.63 -0.20 0.007 0.95

(n = 316) 4 0.05 0.003 0.20 0.14 0.002 0.97 Tyrannidae 5 0.04 0.002 0.47 0.07 0.002 0.98 1 0.18 0.020 0.27 -0.15 0.034 0.68 0.05 2 0.08 0.008 -0.01 0.11 0.006 0.80 3 0.06 0.007 0.64 0.36 0.004 0.89

(n = 89) 4 0.04 0.005 0.60 0.26 0.002 0.92 Muscicapini 5 0.04 0.004 2.43 -1.05 0.001 0.95 1 0.22 0.030 0.68 0.05 0.050 0.66 0.08 2 0.10 0.010 1.23 -0.32 0.010 0.79 3 0.09 0.010 1.21 -0.73 0.007 0.89

(n = 72) 4 0.06 0.007 -0.27 0.06 0.003 0.94 Monarchini 5 0.04 0.004 0.04 -0.20 0.001 0.96

132 Table 3.3. Results of the canonical discriminant analysis on nine morphological variables of Old and New World flycatchers. Values in the table are the coefficients which correspond to the importance of the variable in explaining the between-class variation, the Eigenvalue and the cumulative variance explained by the canonical variate (Cani).

Variable Can1 Can2 Can3 Can4 Total 0.974 0.104 0.146 -0.139 Wing 0.896 0.432 0.102 -0.012 Tail 0.974 -0.080 0.131 -0.168 Tarsus 0.964 -0.138 0.225 0.017 Toe 0.612 0.740 0.267 -0.084 Bill Length 0.952 0.290 0.096 -0.021 Bill Width 1 0.828 0.438 0.350 -0.008 Bill Width 2 0.918 0.393 0.058 -0.008 Bill Depth 0.934 0.187 0.306 -0.005 Eigenvalue 1.84 1.03 0.33 0.05 Cum. Var 0.57 0.88 0.98 1.00

Table 3.4. Lineage centroids for the canonical discriminant analysis. Mon = Monarchini, Ela = Elaeniinae, Flu = Fluvicolinae, Mus = Muscicapini, Tyr = Tyranninae.

Lineage Can1 Can2 Can3 Can4 Mon 0.700 -0.755 -0.557 -0.281 Ela -1.742 -1.769 0.798 0.045 Flu 0.366 -0.213 -0.557 0.390 Mus -1.345 1.232 -0.033 -0.065 Tyr 2.114 0.540 0.791 0.007

133 Table 3.5. Squared Mahalanobis distances (lower triangular matrix) and F statistics (upper triangular matrix) among lineages of Old and New World flycatchers. The bolded value (fluvicolinae to Monarchini) is significant at alpha = 0.0016. All others are significant at alpha < 0.0001. Lineage abbreviations as in Table 4.

Distance Distance From / F value (difference in means) To Mon Ela Flu Mus Tyr Mon - 25.28 3.06 36.42 19.01 Ela 8.94 - 23.41 29.98 51.78 Flu 0.85 8.82 - 21.54 17.66 Mus 8.45 9.86 5.50 - 48.85 Tyr 5.57 20.20 5.58 13.12 -

Table 3.6. Misclassification table of Old and New World flycatcher lineages based upon the nine morphological variables and apriori classification rate proportional to the sample size of the group. The values in column “n” are the number of species in each lineage. The main diagonal (bolded values) are the percent of species reclassified to the lineage. Lineage abbreviations as in Table 4.

Classified Classified Into From Mon Ela Flu Mus Tyr n Mon 55.56 6.94 20.83 8.33 8.33 72 Ela 0 90.24 0 9.76 0 41 Flu 16.39 4.92 57.38 8.2 13.11 61 Mus 1.12 0 7.87 91.01 0 89 Tyr 16.07 0 0 7.14 76.79 56

Table 3.7. Statistical moments of the ellipse analysis. Lineage abbreviations as in Table 4. New = values from grouped New World lineages. Old = values from grouped Old World lineages.

varx vary covxy (E- Lineage n (E-03) (E-03) 03) L1 L2 a12 sed Slope shape seshape ELA 41 3.3 1.3 0.9 0.060 0.031 0.308 0.103 0.32 1.954 0.305 FLU 61 7.5 3.3 2.4 0.093 0.047 0.356 0.082 0.38 1.971 0.252 TYR 56 7.2 0.8 0.75 0.085 0.027 0.109 0.046 0.11 3.196 0.427 NEW 158 7.3 3.2 2.3 0.091 0.047 0.352 0.051 0.38 1.960 0.156 MUS 89 4.0 2.6 0.2 0.063 0.051 0.083 0.234 0.08 1.251 0.133 ARC 72 8.5 5.2 1.4 0.095 0.068 0.239 0.172 0.25 1.387 0.163 OLD 161 7.3 3.7 0.8 0.086 0.059 0.151 0.102 0.15 1.455 0.115 n = Number of species in lineage. varx = Variance in habitat axis. vary = Variance in diet axis. covxy = Covariance of habitat and diet. Li = Standard deviation of axis i of major (1) and minor (2) axes. a12 = Directional cosine of diet on first eigenvector. sed = Standard Error of the directional cosine. -1 Slope between X and Y = tan(sin [a12]). shape = √(first eigenvector / second eigenvector). seshape = Standard Error of shape.

134 135

Table 3.8. T-values of pair wise comparisons in orientations (a12) and shape (√L1/L2) of ellipse parameters among lineages. Lineage abbreviations as in Table 4. P values less than 0.01 are presented in parentheses after the value.

Comparison Orientation Shape NEW-ELA 0.39 0.02 NEW-FLU 0.04 0.04 NEW-TYR* 2.69 (<0.01) 3.39 (<0.01) ELA-FLU 0.37 0.04 FLU-TYR 2.57 (0.01) 2.52 (0.01) TYR-ELA 1.93 (0.057) 2.20 (0.03) OLD-MON 0.46 0.33 OLD-MUS 0.31 1.11 MON-MUS 0.52 0.65 OLD-NEW 1.75 (0.08) 2.61 (<0.01)

Table 4.1. Coefficients and cumulative variance explained in the 1) separate principal component analyses (PCA) and 2) common principal component analysis (CPCA) on elaenia and tody-tyrant lineages. Principal components were computed on corresponding variance-covariance matrices. Values in parentheses after the first coefficients in each ordination are the standard errors (based on 1000 bootstrap runs). These standard errors were used to analyze multivariate allometry in the corresponding variable. Bold values indicate significant allometry (positive or negative) with respect to an expected value of 0.4081 (see text for explanation).

Elaenia PCA Tody-tyrants PCA CPCA VARIABLE PC1 PC2 PC3 PC1 PC2 PC3 CPC1 CPC2 CPC3 wing 0.49 (0.022) -0.22 -0.46 0.45 (0.027) -0.22 -0.002 0.47 (0.020) 0.19 -0.30 tail 0.58 (0.061) 0.55 -0.36 0.64 (0.050) 0.62 -0.38 0.62 (0.040) -0.61 -0.34 tarsus 0.21 (0.040) 0.43 0.46 0.13 (0.036) 0.40 0.45 0.15 (0.030) -0.40 0.67 bill length 0.27 (0.044) 0.19 0.52 0.20 (0.025) 0.08 0.80 0.24 (0.030) -0.10 0.50 bill width 0.46 (0.060) -0.48 0.43 0.46 (0.049) -0.59 -0.03 0.45 (0.040) 0.57 0.31 bill depth 0.31 (0.040) -0.45 -0.02 0.35 (0.040) -0.26 0.12 0.34 (0.029) 0.32 -0.03 lineage values

Etot 0.028 0.004 0.003 eigenvalues 0.028 0.0040.003 0.035 0.007 0.002 Ttot 0.0343 0.0068 0.0017

Etot 75.21 86.23 94.39 % cumulative variance 75.79 87.03 95.19 75.49 90.64 95.19 Ttot 75.24 90.34 94.15

136 137 Table 4.2. Estimated t values of sequential pair-wise comparisons between subclade parameters of ellipse shape and orientation. Values in boldface type are significant at p = 0.05. Boldfaced values with an asterisk are significant to maximum of p = 0.01. The multiplicity correction is based on an alpha level of 0.05/5 = 0.01 where 5 is the maximum number of times any one group is included in a pair-wise comparison.

CPC1 vs CPC2 CPC2 vs. CPC3 Comparison Shape Orientation Shape Orientation E-T 0.77 0.01 0.56 3.52* E-e1 2.37 0.99 1.65 2.87* E-e2 0.65 2.46 1.82 1.22 E-e3 0.13 0.22 0.35 0.03 E-e4 1.07 2.94* 0.76 1.99 e1-e2 0.87 2.9* 0.48 0.80 e1-e3 1.54 0.57 1.32 2.05 e1-e4 2.71* 1.94 1.98 0.11 e2-e3 0.46 1.97 1.55 0.93 e2-e4 1.18 0.24 2.15 0.26 e3-e4 0.70 1.70 0.23 1.13 T-t1 0.38 1.32 0.78 0.10 T-t2 2.48 4.02* 1.68 0.59 t1-t2 1.15 1.61 2.05 0.28

138 Table 4.3. Median Nearest Neighbor Distances (NNDs) for each lineage and subclade (see Fig. 1 for group designation). Delta values are taken from the DNA hybridization work of Sibley and Alquist (1990) and represent relative ages of origin for each group. An increasing age of delta value corresponds to an increasing hypothesized age of origin. The NNDs are computed using CPC scores from corresponding axes.

Median NND Group ∆T50H CPCs 1&2 CPCs 2&3 E 5.7 0.189 0.096 e1 5.0 0.171 0.072 e2 4.0 0.197 0.099 e3 4.0 0.143 0.059 e4 4.5 0.159 0.108 T 9.4 0.183 0.097 t1 7.4 0.243 0.111 t2 7.4 0.107 0.082

139 Figure 1.1. A phylogeny of the Passeriformes based upon Sibley and Alquist DNA Hybridization data. Clades pertinent to the project are highlighted with black lines. Tyrannida includes the tyrant flycatchers (Tyrannidae), Corvidea includes the monarch flycatchers (Monarchini) and the Old World flycatchers (Muscicapini) are included in the Muscicapoidea.

I. Acanthisittides

I. Eurylaimides P. Funariida P. Thamnophilida P. Tyrannida S. Ptilonorhynchoidea

Passeriformes S. Meliphagoidea S. Corvoidea S. Muscicapoidea S. Sylvoidea S. Passeroidea

140 Figure 2.1. The phylogeny based on DNA hybridization data (Sibley and Alquist 1990) of the flycatcher species used in the study of concordance in ecomorphological relationships.

Tyrannus verticalis (Western Kingbird) Tyrannus vociferans (Cassin’s Kingbird) Tyrannus tyrannus (Eastern Kingbird) Myiodynastes luteiventris (Sulpher-bellied Flycatcher) Myiarchus cinearascens (Ash-throated Flycatcher) Myiarchus tuberculifer (Dusky-capped Flycatcher) Sayornis nigricans (Black Phoebe) Sayornis phoebe (Eastern Phoebe) Contopus virens (Eastern Wood Pewee) Contopus pertinax (Greater Pewee) Contopus sordidulus (Western Wood Pewee) Empidonax virescens (Acadian Flycatcher) Empidonax fulvifrons (Buff-breasted Flycatcher) Empidonax traillii (Willow Flycatcher) Pyrocephalus rubinus (Vermillion Flycatcher) Batis Species Terpsiphone viridis (Paradise Flycatcher) Bradornis mariquensis (Marico Flycatcher) Melaenornis pammelina (Southern Black Flycatcher) African Taxa Sigelus silens (Fiscal Flycatcher) except M. pictus Muscicapa striata (Spotted Flycatcher) Myioparus plumbeus (Lead-colored Flycatcher) Myioborus pictus (Painted Redstart)

141 Figure 2.2. Joint scatterplot of the species scores in search foraging of New World flycatchers. The scores are derived from a correspondence analysis of the cross- classified enumerated percentages of two categorical variables. O = open habitat, C = Closed habitat. 1-5 is a gradient between low perches (1) and high (5). Species codes are as in Table 3. Dotted lines are drawn through the origins.

2 O1 C1

1 sn mcC2 O2 et C3 O4 sp pr mp 0 C4 tvo ef mt ev tve ml cs cv Search 2 O3 tt C5 -1 O5cp

-2 -2 -1 0 1 2 Search 1

142 Figure 2.3. Joint scatterplot of the species scores in attack foraging of Old and New World flycatchers. The scores are derived from a correspondence analysis of the enumerated percentages of foraging mode. H = hoverglean, S = standpick, F = flycatch, P = pounce. Species codes are as in Table 3. Dotted lines are drawn through the origins.

2

HMPb S

1 BAT TV mt BM

ev

Attack 2 etmp P ef SS 0 MP sp cv sn MS mcprml cs tvo tttve Fcp -1 -10123 Attack 1

143 Figure 2.4a & b. Scatterplot of the average species scores of Old and New World flycatchers from a principal components analysis on log10 transformed morphological variables. Species codes are as in Table 3. Dotted lines are drawn through the origins.

a. long tail, tarsus, toe F BF 0.2 BM

mp MPb 0.1 snTV mc sp 0.0 MSpr ef et mt tt tvetvo Morph 2 ev cp -0.1 cvcs wide, BAT ml deep bill -0.6 -0.4 -0.2 0.0 0.2 0.4 0.6 small large Morph 1

b. long 0.1 tarsus, toe F etmt mp ml ev ef BM MPb pr spmc 0.0

tvott sn BF tve BATcscvcp

-0.1 TV Morph 3 MS long wing -0.1 0.0 0.1 0.2 wide, long tail, deep bill Morph 2 tarsus, toe

144 Figure 2.5. Scatterplot predicting search behavior from morphology in the New World flycatchers. Along the Y axis, the positive end (“1”) is associated with low perch heights and the negative end (“5”) is associated with high perch heights. The upper values with species codes as the symbols (as in Table 3) are from the canonical correlation analysis on the species’ mean scores from the first three PCA (morphology) and first three CA (search behavior) axes. To distinguish between upper and lower distributions, I added a constant (3) to the upper search scores. The lower plot with open circles depicts the relationship between the first pair of PLS variables using phylogenetically independent contrasts computed from the morphology and search behavior scores using the first pair of PLS scores. The PLS regression was run through the origin. Both regressions are significant (see text).

6 NA Flycatchers R2 = 0.7* 1 5 sn mc 4 et mp pr 3 ml efmtsp cs tvo ev 2 tvett cv cp 1

Search CCA1 2 0 R = 0.5*

5 -1 -2 -3 -2 -1 0 1 2 3 Large bills Small bills Short Legs Morphology CCA1 Longer Legs

145 Figure 2.6 a&b. Scatterplot of the canonical correlation analysis of attack behavior and morphology. The plots depict the linear relationship between morphology and attack behavior. The top figure (a) is comprised of the first canonical axis constructed from principal component scores on morphology and scores from a correspondence analysis on attack foraging. Flycatcher communities are separated into Old (o) and New (n) World species. Phylogenetically independent contrasts were computed from the PCA and CA scores and their relationship in canonical space is presented in figure b. Least squares regression lines (dashed for the Old World species and solid for the New World species) are drawn through the origin for the contrast plot. Accounting for phylogeny seems to have little effect on the prediction of foraging from morphology. The asterisk denotes the contrast between Muscicapa striata and Myioparus plumbeus. The cross denotes the Root contrast.

a. 3 BM

2

MPSS 1 MPbmp TV 0 BAT MS sp

Attack CCA 1 tvo mt pr sn ml evttytve et ef -1 cscbcv mc

-2 -1 0 1 2 Morphology CCA 1

b. 3 o

n 2

1 n on o nn n nn n o 0 n n Attack Contrasts CCA 1 o

0 1 2 3 Morphology Contrasts CCA 1

146 Figure 2.7. Attack rate (Foraging Velocity) plotted against the first morphological PC. The first PC is a size axis; large species tend toward the right, small species are on the left. The Painted Redstart and combined Batises (crosses) were not included in the regression analysis. Attack rate is measured as a grand mean for each species. The vertical lines are the negative standard errors. Note the break in the Y axis with respect to the redstart attack rate in comparison to the flycatchers.

0.50

0.45 mp

ef BAT

0.10 mt ev cv et sn mc

pr cs tt

Foraging Velocity (attempts/second) sp MP BM ml cp SS tve tvo 0.00

-0.6 -0.4 -0.2 0.0 0.2 0.4 0.6 Morphology PC1 (Size)

147 Figure 2.8 . Percent of foraging maneuvers of Old and New World species that are comprised of flycatching (Black portion of bar) and of other maneuvers (white portion of bar). The dashed line is drawn at approximately 50% mark for reference. The solid line in the species list separates Old (below) and New (above) World species.

148 Figure 3.1. Scatterplot of the first and second PC axes from separate covariance matrices of three flycatcher lineages. The first axis in all cases describes the overall variation in size within each lineage. The second axis describes shape variation within each group, but unlike the first axis the variables that are important in explaining the variation within each lineage is different for each.

0 Tyrannidae n = 316 Muscicapini n = 89 0 Monarchini n = 72 Shape Axis (PCs2) (PCs2) Axis Shape Lineage Dependent 0

-1 -0.5 0 0.5 1 Common Size Axis (PCs1)

149 Figure 3.2. Frequency distributions of species scores along the first three principal components for each group. Groups: Tyr = Tyrannidae, Mus = Muscicapini, Mon = Monarchini. The species are divided into 15 bins of equal size (range / 15).

Tyr Mus Mon 50 40 PC1 30 20 10 0 50 40 PC2 30 20 10 0

Frequency 100 80 PC3 60 40 20 0 Bin

150 Figure 3.3. Sign, magnitude and similarity of each variable within each eigenvector of the separate principal components analyses of three flycatcher lineages. The sham variable is the alternating minimum and maximum value for each principal component across all variables for each lineage. If lineages are similar in morphology (convergence) across PCs with respect to magnitude and sign of each variablem, patterns within each variable should show parallel change from one PC to the next as seen in the variable. The first PC is an overall size axis. Hence, the lineage points for that variable are positive and clustered tightly together. The second and remaining PCs are shape variables. These axes show much less similarity across the lineages. For example, Bill Depth shows similarities between Muscicapini (short dashed lines) and Monarchini (alternating dot and dash lines) but in the tyrants (solid lines), bill depth is negatively loaded. These differences are more apparent, for example, in tarsus length. The variable within the second PC is loads highly and on the positive side of the origin for the Monarchini and Tyrannidae, but loads highly and on the negative side of the origin for the Muscicapini.

151 Figure 3.4. The relative rates of overall body size and shape evolution. These were obtained by dividing the standard deviations of the species scores along the first (size = black bars) and second (shape = white bars) PCs by the nodal delta value for each group from Sibley and Alquist (1990).

0.05 Size Shape

0.04

0.03

0.02 Rates of Diversification of Rates 0.01

0.00 Tyrannidae Muscicapini Monarchini

152 Figure 3.5. The morphological space occupied by the five lineages of flycatchers in canonical discriminant space. “Elaeniinae = filled circles, “fluvicolinae” = open squares, “tyranninae” = closed triangles, Muscicapini = 4s, Monarchini = 5s. The first axis is an overall size axis, so species towards the right (i.e. “tyranninae”) are relatively large, and species towards the left of the axis (i.e. Muscicapini and “elaeniinae”) are relatively small. Along canonical axis two species in the upper part of the have relatively long toe and wing and a wide bill but a short tarsus. Scores at the lower end of the axis are depicting species with opposite features.

-4 -2 0 2 4

4 4 Toe 44 4 Wing 4 4 4 4 4 4 44 4 4 4 4 4 4 2 4 44 4 4 4 4 4 4 44 4 4 44 44544 444 444 5 4 44 4 444 5 44 44 55 4 4 4 54 4 5 5 44 444 4 444444 55 5 4 5 5 5 5 5 0 4 45 4 55 5 5 55 4 5 54 5 4 4 5 5 4 44 5 55 555 Can2 5 555 5 5 4 5 55 5 5 5 5 55 5 5 5 5 5 55 5 55 55 5 5 55 -2 5 5 5 55 Tarsus 5 -4 5

MUS Can1 (Size) TYR FLU MON ELA

153 Figure 3.6. The morphological space occupied by the five lineages of flycatchers in canonical discriminant space. Darkened circles are New World flycatchers while open squares are Old World flycatchers. Ellipses are drawn around each group that contains species within one standard deviation of the centroid.

4 3 2 1

2 0

n

a

C -1 -2 -3 -4 -5 -5 -4 -3 -2 -1 0 1 2 3 4 5 Can1 (Size)

154 Figure 3.7. UPGMA cluster analysis on size-free generic mean morphological traits. The first letter of the genus corresponds to the lineage: e = Elaeniinae, f = fluvicolinae, t = tyranninae, U = Muscicapini, O = Monarchini. The genus is represented by the first seven letters of the taxonomic name of the genus and the symbol depicts whether the species within each genus generally utilize Open (filled circle) or Closed (open triangle) habitats. The tick mark represents a cluster largely made up of non-phylogenetically grouping genera comprised of species that primarily utilize closed habitats.

155 Figure 3.8. The evolutionary rates and changes in tempo of diversification in flycatcher subclades. I divided the standard deviation of the major and minor axes of the ellipse of each Old and New World lineage and their combined total values by their estimated relative age of origin (the nodal delta values from Sibley and Alquist (1990)).

L2/age L1/age 0.02

0.01 / Relative age of orgin) 2 Rate of Diversification and L L and 1

(L (L 0 FLU ELA OLD TYR MUS NEW MON

156 Figure 4.1. Lineage specific patterns of differsification in bivariate space. Along the axes are plotted values for two traits. Nintey-five percent confidence ellipses are drawn around species within monophyletic subclades. The shape and orientation of the bivariate ellipses give quantitative measures of mode and tempo of trait evolution. The lineages can be either independent clades (e.g. A = muscicapidae B = tyrannidae) or hierarchical (e.g. c = mionectinae and d = fluvicolinae within B = tyrannidae). In this figure, both the orientation and shape of the bivariate ellipses give evidence as to the mode and tempo of evolution of traits 1 and 2. For example, lineage B has diversified at a more rapid tempo than A along the “trait 2” axis when compared lineage A and at a more rapid tempo than it has along the “trait 1” axis. However, the subclade disparities within B are oriented along a “Trait 1” axis. In addition, compared to B (the mother lineage), subclades c and d occupy a smaller volume and have different orientations as well as different shapes. One may infer from this that the early diversification in lineage B was along a “trait 2” axis while later diversification (the subclades) occurred along a “trait 1” axis. For further review, see Derrickson and Ricklefs (1988) and Ricklefs and Nealen (1989).

c A B Trait 2 d

Trait 1 (PC1)

157 Figure 4.2. Synthesized phylogeny from Sibley and Alquist (1990) DNA and Lanyon (1988a; 1988b) morphological data. Number of species is in parentheses for each genus and subclade. T = tody-tyrant lineage; E = elaenia lineage. Note: Atalotriccus pilaris and Lophotriccus spp. are combined; Cnipodectes subbrunneus is only included in Ttot.

Genera Subclades

Rhynchocyclus Tolmomyias Onychorhynchus T1 Platyrinchus Cnipodectes Todirostrum Ttot Poecilotriccus Myiornis T1 Hemitriccus Oncostoma Lophotriccus Elaenia Myiopagis Tyranniscus E1 Mecocerculus 1 Ornition Camptostoma Mecocerculus 2 E2 Suiriri Mecocerculus 3 Etot Capsiempis Phaeomyias E4 Nesotriccus E3 Serpophaga Uromyias Anairetes Pseudocolopteryx Polystictus

158 Figure 4.3. Standard errors about bootstrapped (n=1000) coefficients plotted against the eigenvectors of a common principal components analysis on six log-transformed morphological traits of elaenia and tody-tyrant flycatcher lineages. The individual lines correspond to the six morphological variables. Notice that the standard errors dramatically increase after the third eigenvector.

0.6

0.5

0.4

0.3

0.2 SEs for Variables SEs 0.1

0 123456 CPC Bootstrapped Eigenvectors

159 Figure 4.4. Scatterplot of elaenia along the first two common principal component axes. The ellipses were computed and drawn around species members in each subclade. Solid lines are used around the subclades while the whole-clade disparity is characterized by a dashed line. The ellipse around the sublade e4, which consists of e2, e3, Pseudocolopteryx and Polystictus was omitted for clarity. The ellipse parameters broken down by subclade are listed in Appendix IV.

0.8 Short tail Wide bill e1 e2 e3

0.6

0.4 CPC2

Long tail Narrow bill 0.2 2.4 2.6 2.8 3.0 3.2 3.4 CPC1 (size)

160 Figure 4.5. Scatterplot of tody-tyrants along the first two common principal component axes. Note: Myiornis ecaudatus (CPC1 = 2.37, CPC2 = -0.376) was omitted from the graph for correspondence of axes in other figures. The asterisks by the triangle in the left- hand side and by the four circles in the right-hand side of this figure are to note the extreme positions of the two Myiornis species, and the four right-hand species (Rhynchocyclus spp. and Cnipodectes subrunneus) respectively. The dashed line represents the confidence ellipse for the entire clade. See text for explanation.

0.8 Short tail Wide bill

0.6 *

*

0.4 CPC2

Long tail t1 Narrow t2 bill 0.2 2.42.62.83.03.23.4 CPC1 (Size)

161 Figure 4.6. Scatterplot of elaenia along the second and third common principal component axes. Lines as in Figure 4.4.

LongLon g0.7 billbill Long legsLon g legs 0.6

0.5

CPC3 e1 e2 e3 0.4 ShortShor bill Shortt bill Shorlegs t 0.20.30.40.50.60.70.8Long tail Short LongNarrow tail Short tail tail Narrow bill CPC Wide bill

162 Figure 4.7. Scatterplot of tody-tyrants along the second and third common principal component axes. Lines as in Figure 4.5.

Long bill 0.7 Long legs

0.6

0.5 CPC3

t1 Short 0.4 bill t2 Short legs

0.2 0.3 0.4 0.5 0.6 0.7 0.8 Long tail Short tail Narrow bill CPC2 Wide bill

163 Figure 4.8. Total lineage morphological disparity in of elaenia (E) and tody-tyrant (T) flycatchers based on a characterization of the morphological volume from a common principal components analysis. CPC1 is the first axis explaining a majority of the variance in both lineages and is considered an overall size disparity axis. CPC2 is a shape axis and explains the next largest proportion of variance in the lineage data sets. Species within each lineage with short tails and wide bills are at the high end of the axis while species in both lineages with long tails and narrow bills score at the lower end of the axis.

CPC1 CPC2

3.3 -0.34

3.0 -0.51

2.7 -0.68

2.4 -0.85 E T E T

Appendix I. Species, sample sizes and morphological means and standard deviations for tyrant flycatchers. An asterisk denotes the species used in the following analysis (Appendix II). Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 1 Agriornis andicola * 5 260.8 11.2 134.4 4.5 113.8 4.1 38.3 2.6 29.7 1 2 Agriornis livida * 10 263.8 18.1 129.7 4.7 116.8 2.7 37 2.2 29.4 1.8 3 Agriornis microptera * 7 239.9 19.9 115.9 6.3 111 11.5 32.9 1 24.8 0.9 4 Agriornis montana * 7 236.1 12 134.7 6.4 109 4.2 33.1 1.4 25.2 1.4 5 Agriornis murina * 7 183.1 8.2 99.1 3.2 82.4 2.9 27.5 1 19.7 0.9 6 Alectrurus risora 5 224.8 - 73 3.3 141 - 21.9 0.6 19.3 2.1 7 Alectrurus tricolor 1 104 - 64 - 35 - 18.9 - 18.7 - 8 Anairetes alpinus * 4 131 1.8 62.8 1.5 69.8 1.7 20.7 0.6 14.8 0.6 9 Anairetes fernandezianus* 0 ------10 Anairetes flavirostris * 10 102.6 4.7 48.8 2.7 52.6 6.5 17.9 0.7 11.8 0.6 11 Anairetes nigrocristatus* 0 ------12 Anairetes parulus 11 104.6 4 47 1.8 51.7 2.2 18.5 1.2 12.1 0.7 13 Anairetes reguloides 10 124.8 7.5 56.8 2.5 66.6 4.1 20.5 1.6 14.1 0.5 14 Aphanotriccus audax 10 125.3 5.9 60.3 2.9 57.5 4.1 14.3 0.7 12.8 0.8 15 Aphanotriccus capitalis 2 116.5 9.2 58.5 0.7 57.5 0.7 13.9 0.1 12.7 0.6 16 Arundinicola leucocephala 10 123.5 4.4 62 1.8 47.6 2.3 16.3 0.4 16 1.5 17 Atalotriccus pilaris 8 90.5 3 40.4 1.6 37.6 3.2 15.8 0.7 10.9 0.3 18 Attila bolivianus * 2 186 8.5 96 4.2 90.5 6.4 24.8 0.1 22.8 1.2 19 Attila cinnamomeus * 10 184 3.8 91.4 3.8 87.9 4.1 21.8 0.9 19.5 1.3 20 Attila citriniventris * 6 175 8.6 88.7 2 75.3 2.1 21.9 0.6 19.8 0.9 21 Attila phoenicurus * 2 188.5 2.1 93 1.4 87 0 19.3 0.1 18.6 0.6 22 Attila rufus * 10 196.1 11.8 91.4 4.2 82.5 3.6 24.8 0.8 21.7 1.1 23 Attila spadiceus * 8 181 12 87.6 5.9 77.5 5.9 25.3 1.3 23.4 1.3 24 Attila torridus * 3 218 2.6 95 5.3 90 5.2 25.6 1 22.8 1.3 25 Camptostoma imberbe * 10 105.3 5.6 53.9 3.2 46.2 4.6 13.9 0.6 11.4 0.4 26 Camptostoma obsoletum * 12 106.8 6 54 2.2 43.6 4.7 14.3 0.8 12 0.8 27 Capsiempis flaveola * 10 109.7 8.5 48 2.1 51.2 5.1 16 0.9 11.1 1.3 28 Casiornis fusca 2 168 2.8 77 1.4 81 5.7 19 0.2 16.3 0.8 29 Casiornis rufa 10 170.3 8.1 83.4 3.3 83.4 4.7 19.8 0.6 15.7 1.2 164 30 Cnemarchus erythropygius 10 227.2 6.3 142.3 2.9 112.7 2.9 28.8 1.4 24.3 1.7

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 1 Agriornis andicola * 5 33.3 1.4 17.7 1.4 7.5 0.3 8.6 0.4 2 Agriornis livida * 10 35 1.2 18.2 1.2 7.6 0.4 8.1 0.5 3 Agriornis microptera * 7 31.4 1.2 17 1.3 7.3 0.3 7.6 0.5 4 Agriornis montana * 7 30.8 1.3 14.5 0.8 6.3 0.4 6.2 0.3 5 Agriornis murina * 7 22.3 0.7 11.5 1.3 4.9 0.3 4.7 0.6 6 Alectrurus risora 5 18.7 0.8 12 0.9 7.2 0.9 5.4 0.3 7 Alectrurus tricolor 1 16.8 - 9.3 - 6.6 - 4.8 - 8 Anairetes alpinus * 4 12.7 0.9 6.7 0.6 3.1 0.6 3.2 0.2 9 Anairetes fernandezianus* 0 ------10 Anairetes flavirostris * 10 11.6 0.5 6.4 0.4 2.6 0.2 2.8 0.2 11 Anairetes nigrocristatus* 0 ------12 Anairetes parulus 11 12.3 0.5 6.1 0.5 2.6 0.2 2.7 0.4 13 Anairetes reguloides 10 13.5 0.7 6.8 0.5 3 0.3 3.4 0.4 14 Aphanotriccus audax 10 14.8 0.4 9 0.4 6.3 0.2 3.8 0.4 15 Aphanotriccus capitalis 2 14 0.4 8.4 0.3 5.9 0.2 3.9 0 16 Arundinicola leucocephala 10 18.1 1.1 9.5 0.7 5.4 0.3 4.3 0.3 17 Atalotriccus pilaris 8 12.4 0.8 6.5 0.3 3.8 0.4 2.9 0.3 18 Attila bolivianus * 2 27 0.3 14.9 0 8 0.1 6.3 0.2 19 Attila cinnamomeus * 10 23.2 1.2 14.5 1 7.2 0.8 6.2 0.4 20 Attila citriniventris * 6 24.5 0.8 15.1 0.9 7.5 0.8 6.6 0.3 21 Attila phoenicurus * 2 21.8 0.1 12.9 1.4 6.5 0.3 5.6 0.1 22 Attila rufus * 10 29.3 1 15.4 0.9 7.9 0.4 6.6 0.3 23 Attila spadiceus * 8 27.4 1.9 15.6 1.3 7.7 0.4 6.7 0.6 24 Attila torridus * 3 27.7 1.6 15.8 2 8.8 0.1 6.8 0.3 25 Camptostoma imberbe * 10 11.4 1.9 6.6 0.6 3 0.2 3.3 0.3 26 Camptostoma obsoletum * 12 10.7 0.5 6.9 0.6 3.3 0.2 3.4 0.3 27 Capsiempis flaveola * 10 12.2 0.5 7.1 0.9 3.6 0.4 3.6 0.2 28 Casiornis fusca 2 19.2 0.6 10.3 1.5 5.7 0.7 4.7 0.6 29 Casiornis rufa 10 19.6 0.7 11.7 1.1 6.3 0.5 5.2 0.4 30 Cnemarchus erythropygius 10 23.2 0.9 13.2 1.5 5.9 0.4 6.1 0.4

165

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 31 Cnemotriccus fuscatus 10 131.9 4.4 65.4 3.4 67.1 4.5 16.9 1.3 14.2 1.2 32 Cnipodectes subbrunneus 10 171 10.2 87.7 6.1 84.4 6.3 17.5 0.8 16.6 1.1 33 Colonia colonus 10 124.3 8.6 71.7 5.5 58.8 3.8 13.7 0.8 13.7 1 34 Colorhamphus parvirostris 9 126.1 5.1 62.8 4.3 59.7 4.6 17.7 1.3 14.4 1 35 Conopias albovittata 7 150.7 6 81.6 3.6 67.4 4.4 17.6 0.5 19.1 1.1 36 Conopias cinchoneti 2 158 8.5 85.5 0.7 74.5 0.7 16.9 0.2 17.5 2 37 Conopias parva 3 152.3 6.4 77.7 3.5 68.3 1.5 15.9 1 19.2 1 38 Conopias trivirgata 1 133 - 69 - 63 - 16.3 - 16.9 - 39 Contopus borealis * 11 176.6 10.6 105.2 4.6 73.1 4.1 14.4 0.6 16 1 40 Contopus caribaeus * 11 144.5 5.3 73.2 2.2 70.1 3.2 15.2 0.6 12.4 0.7 41 Contopus cinereus * 10 132.5 6.4 71.2 2.7 64.2 3.5 12.7 0.5 12 0.6 42 Contopus fumigatus * 1 156 - 92 - 71 - 14.6 - 13.2 - 43 Contopus hispanolensis * 8 135.9 3.5 75.6 3.5 69.9 2.6 14.4 0.7 12.4 0.8 44 Contopus latirostris * 7 131.7 3.9 65.9 2.5 65.7 2.7 15 0.5 11.8 0.4 45 Contopus lugubris * 10 162.1 7.2 92.1 2.2 80.4 2.3 15.3 0.4 14.3 0.5 46 Contopus nigrescens * 4 120.8 4.5 65.3 2.4 60.3 2.2 11.1 0.4 10.3 0.6 47 Contopus ochraceus * 4 158.3 4.2 88.5 3.1 75.8 2.2 15.4 0.5 14 0.9 48 Contopus pertinax * 11 171.4 6.6 101.7 4.8 86.3 4.1 16 0.9 14.8 1.1 49 Contopus sordidulus * 10 145.5 4.3 84.8 2.4 69.3 2.2 13.3 1.8 12.5 0.3 50 Contopus virens * 11 147 6.8 83.1 3.4 68.3 3.1 12.8 0.4 12.7 0.8 51 Corythopis delalandi 9 125 8.3 62.1 3.2 55 4.4 23.2 0.9 17.6 0.9 52 Corythopis torquata 16 128.3 6.5 65.8 4.7 55.6 4.9 24.8 1 17.7 0.9 53 Culicivora caudacuta 9 99.2 4.2 40.7 2.2 47.4 3 14.4 1.5 12.1 0.5 54 Deltarhynchus flammulatus 7 156.1 4.4 73.7 2.8 74 7.4 17.7 0.5 14.7 0.7 55 Elaenia albiceps * 14 140.4 5 75.4 2.7 68.3 4.4 18.5 1.1 14.5 1 56 Elaenia chiriquensis * 13 140 5.1 75.1 2.9 67.9 4.2 16.8 1.5 13.4 1.1 57 Elaenia cristata * 13 133.8 5.4 67.2 2.6 63.2 2.8 17.9 1 13.2 0.7 58 Elaenia dayi* 0 ------59 Elaenia fallax * 10 144.7 2.2 66.6 3.4 70.9 2.6 18.3 1 13.8 0.8 60 Elaenia flavogaster * 11 156.3 7 77.6 3.9 72.6 5.4 18.6 1.1 15.3 0.8

166

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 31 Cnemotriccus fuscatus 10 16 0.8 8.6 0.8 5.9 0.7 4.2 0.2 32 Cnipodectes subbrunneus 10 19.3 1.3 12.6 0.8 7.5 0.8 5.1 0.4 33 Colonia colonus 10 12.2 0.6 10.3 0.5 5.3 0.5 3.3 0.3 34 Colorhamphus parvirostris 9 12.7 1.1 6.9 0.8 4.2 0.4 3.2 0.3 35 Conopias albovittata 7 20.5 0.8 12.7 0.9 7.8 0.5 5.9 0.2 36 Conopias cinchoneti 2 20 0.4 12.1 0.4 6.4 0.6 5.7 0.7 37 Conopias parva 3 21.2 0.8 12.6 0.1 8.6 0.6 6.2 0.5 38 Conopias trivirgata 1 16.5 - 9.2 - 5.7 - 4.4 - 39 Contopus borealis * 11 22.4 1.1 13.6 0.6 8.7 0.4 6.8 0.5 40 Contopus caribaeus * 11 20 0.8 8.7 0.7 7.4 0.6 4.3 0.5 41 Contopus cinereus * 10 16.8 0.8 8.8 0.6 6.6 0.3 4 0.5 42 Contopus fumigatus * 1 19.7 - 13.1 - - - 5.1 - 43 Contopus hispanolensis * 8 18 0.6 7.9 0.6 6.3 0.3 3.7 0.3 44 Contopus latirostris * 7 17 0.5 8.5 0.8 6.9 0.3 3.6 0.3 45 Contopus lugubris * 10 21.1 1.7 10.7 0.4 7.7 0.4 5.2 0.3 46 Contopus nigrescens * 4 14.1 0.3 8.3 0.6 5.7 0.4 3.2 0.2 47 Contopus ochraceus * 4 20.9 1 10.9 0.7 8 0.6 5.1 0.2 48 Contopus pertinax * 11 21.5 1.3 11.8 0.7 8 0.5 6.1 0.4 49 Contopus sordidulus * 10 17.7 0.9 9.8 0.4 6.8 0.3 5 0.5 50 Contopus virens * 11 17.7 1 9.8 0.5 6.3 0.4 4.8 0.3 51 Corythopis delalandi 9 16.6 0.9 8.4 0.6 4 0.2 3.7 0.2 52 Corythopis torquata 16 16.7 0.7 8.5 0.5 4.2 0.3 4.1 0.3 53 Culicivora caudacuta 9 11.3 0.3 6.8 0.6 4.1 0.3 3.5 0.3 54 Deltarhynchus flammulatus 7 18.4 1 13.1 0.5 7.6 0.4 4.7 0.3 55 Elaenia albiceps * 14 13.7 0.6 8.9 0.7 - - 3.9 0.3 56 Elaenia chiriquensis * 13 13.6 0.7 9.4 1.3 4.4 0.4 3.7 0.2 57 Elaenia cristata * 13 14.2 0.6 9 1 4.8 0.4 4.4 0.5 58 Elaenia dayi* 0 ------59 Elaenia fallax * 10 12.8 0.3 7.4 0.9 3.8 0.3 3.5 0.3 60 Elaenia flavogaster * 11 14.3 0.7 11.3 1 - - 4.1 0.2

167

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 61 Elaenia frantzii * 14 148.6 7.2 78.4 3.9 73.1 4.4 17.3 1.1 14.4 0.8 62 Elaenia gigas * 7 172.1 5 90.3 3.8 82.6 3.5 18.5 0.6 16.6 2.4 63 Elaenia martinica * 11 153.3 7.7 77.9 3.5 73.9 4.5 20.2 1.3 15.1 0.9 64 Elaenia mesoleuca * 11 148.5 5.1 75.1 2.9 69.2 3.3 16.7 0.8 13.8 1 65 Elaenia obscura * 12 163.4 11 81.9 4.6 82.7 4.9 19.6 1.3 15.7 0.8 66 Elaenia pallatangae * 10 139.8 4.1 73.9 2.3 70.9 2.1 16.9 0.6 14.1 0.7 67 Elaenia parvirostris * 11 134.4 3.6 69.4 2.4 65.3 3.4 17.5 0.8 14 0.9 68 Elaenia pelzelni * 11 171.6 8.3 89.5 3.9 82.2 4.9 21.3 0.7 17 0.7 69 Elaenia ridleyana* 0 ------70 Elaenia ruficeps * 11 112.2 5.7 57.2 3.3 48.6 4.2 22.3 1.1 16.5 1 71 Elaenia spectabilis 11 171.3 7.4 88.1 3.8 85.2 5.9 21 1.1 15.5 1.5 72 Elaenia strepera 2 149.5 0.7 82 1.4 70.5 2.1 16.4 1.5 14 0.7 73 Empidonax affinis * 10 133 4.7 71.6 3.8 65.5 4.7 16.4 0.5 12.7 0.7 74 Empidonax albigularis * 9 120.3 6.8 62.1 3.8 56.7 3.4 16.2 0.3 14.1 0.9 75 Empidonax alnorum * 1 136 - 68 - 52 - 15.8 - 10.9 - 76 Empidonax atriceps * 10 115.9 4.4 59.5 2.5 55.3 2.5 15.5 0.8 13.2 0.8 77 Empidonax difficilis * 19 131.7 5.5 67.7 3.9 63.1 5.5 16.2 0.7 13.2 1 78 Empidonax flavescens * 10 124.5 6.3 67 3.6 58 2.6 16.3 0.4 13.3 0.6 79 Empidonax flaviventris * 10 124.2 5.2 64.7 1.7 54 2.4 16 0.6 12.8 0.9 80 Empidonax fulvifrons * 10 107.8 4.2 58.5 2.4 50.5 2.4 13.7 1.1 11.8 0.5 81 Empidonax hammondii * 13 125.9 7.5 68.4 3.1 59.8 3.9 15.7 0.7 12.8 0.9 82 Empidonax minimus * 10 122.6 5.3 62.2 1.5 58.5 3.1 16.2 1.1 12.6 0.9 83 Empidonax oberholseri * 10 135.5 7.3 67.7 3 63.5 4 17.3 0.8 13.6 0.7 84 Empidonax occidentalis * 2 122.5 0.7 62.5 2.1 59.5 3.5 17 1.3 13 0.2 85 Empidonax traillii * 10 131 4.8 69.3 3.6 60.2 2.8 16 0.9 13.9 0.6 86 Empidonax virescens * 10 134.8 5.6 71.8 3.6 62 4.2 15.3 0.6 13.5 0.7 87 Empidonax wrightii * 18 132.9 6.3 71.2 2.7 63.9 3.3 17.9 0.8 13.8 0.8 88 Empidonomus varius 8 176.5 5.6 97.4 4.3 84.6 4.4 16.3 0.6 15.7 0.7 89 Euscarthmus meloryphus * 11 100.1 3.2 45.9 1.7 44.7 2.2 19.5 0.9 13.6 0.7 90 Euscarthmus rufomarginatus * 1 96 - 41 - 50 - 17.5 - 13 -

168

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 61 Elaenia frantzii * 14 14 0.7 8.8 1.1 4.5 0.4 4 0.3 62 Elaenia gigas * 7 15.1 0.7 11.7 0.6 5.5 0.2 4.8 0.2 63 Elaenia martinica * 11 15.2 1.2 11.4 1 - - 4.3 0.3 64 Elaenia mesoleuca * 11 13.8 0.6 9.3 1 - - 3.8 0.3 65 Elaenia obscura * 12 14.9 0.6 9.9 0.9 4.5 0.3 4.6 0.5 66 Elaenia pallatangae * 10 13.7 0.5 7.8 0.5 4.1 0.3 3.9 0.4 67 Elaenia parvirostris * 11 13.4 0.7 8.6 0.6 - - 3.5 0.2 68 Elaenia pelzelni * 11 16.7 0.8 12.2 0.9 - - 4.5 0.4 69 Elaenia ridleyana* 0 ------70 Elaenia ruficeps * 11 13.3 0.4 7.6 0.8 3.8 0.2 3.5 0.2 71 Elaenia spectabilis 11 15.6 0.8 10.8 0.5 - - 4.3 0.3 72 Elaenia strepera 2 13.3 0.8 10.6 1 - - 3.8 0.1 73 Empidonax affinis * 10 14.8 0.5 7.7 0.7 4.8 0.3 3.7 0.4 74 Empidonax albigularis * 9 15.2 0.8 7.5 0.5 5.9 0.3 3.5 0.4 75 Empidonax alnorum * 1 15.4 - 9.5 - - - 3.8 - 76 Empidonax atriceps * 10 13.1 0.6 6.9 0.7 4.7 0.4 3.4 0.4 77 Empidonax difficilis * 19 15.5 0.7 8.2 0.7 5.8 0.3 3.8 0.3 78 Empidonax flavescens * 10 14.9 0.8 8.1 0.4 5.5 0.6 3.9 0.4 79 Empidonax flaviventris * 10 14.5 0.6 8.4 0.3 5.6 0.3 3.8 0.2 80 Empidonax fulvifrons * 10 13 0.5 7.1 0.5 4.5 0.5 3.4 0.4 81 Empidonax hammondii * 13 14.3 1.1 7.2 0.5 4.6 0.2 3.7 0.3 82 Empidonax minimus * 10 14.5 0.5 8 0.8 5.2 0.4 3.8 0.3 83 Empidonax oberholseri * 10 14.9 1.1 8.3 0.5 5 0.3 3.9 0.4 84 Empidonax occidentalis * 2 14.9 0.2 8 0.4 5.3 0.3 4.2 0.3 85 Empidonax traillii * 10 16 0.5 8.4 0.3 5.8 0.3 4.1 0.3 86 Empidonax virescens * 10 16.7 0.7 9.1 0.6 6.3 0.2 4 0.3 87 Empidonax wrightii * 18 16.5 0.6 8.4 1 5.2 0.4 3.9 0.2 88 Empidonomus varius 8 17.5 0.4 12.1 1 7.1 0.9 5.3 0.5 89 Euscarthmus meloryphus * 11 12.6 0.7 6.6 0.6 3.1 0.2 3.1 0.3 90 Euscarthmus rufomarginatus * 1 12.9 - 7.5 - - - 3.3 -

169

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 91 Fluvicola albiventer * 10 130.3 5.9 66.6 4.8 57.6 4.8 20.7 0.6 17.3 1.1 92 Fluvicola nengeta * 7 138.4 8.9 73.9 5.3 63.9 6.1 23 1.3 17.9 0.8 93 Fluvicola pica * 10 126.1 5.2 64 4 53.3 3.2 20.5 0.7 17.1 1.1 Griseotyrannus 94 aurantioatrocristatus 9 165.9 2.6 95.7 3.8 83.1 3 15 0.8 15.5 1.3 95 Gubernetes yetapa * 4 389.3 13.6 122.3 4.5 267.3 16.1 27 1.8 25.4 2 96 Hemitriccus aenigma 6 84.5 3.7 46.5 1.6 36.8 3.9 14.3 0.3 11.1 0.5 97 Hemitriccus cinnamomeipectus 6 103.7 5.6 49.2 2.5 48.2 2.2 18.2 0.7 13 0.4 98 Hemitriccus diops 8 112.5 5.2 51.9 3.5 51.8 4.4 18.1 0.3 13.7 0.5 99 Hemitriccus flammulatus 31 104.4 6.2 53.3 3.8 47.1 4.6 17.6 0.9 12.6 0.7 100 Hemitriccus furcatus 1 86 - 39 - 33 - 14.6 - 8.8 - 101 Hemitriccus granadensis 23 97.6 3.9 46.2 1.9 43.1 2.5 16 0.7 12.4 0.8 102 Hemitriccus inornatus 0 ------103 Hemitriccus iohannis 5 102.6 8 48.6 3.9 41.6 3.6 16.4 2.1 12.1 0.8 104 Hemitriccus josephinae 0 ------105 Hemitriccus kaempferi 0 ------106 Hemitriccus margaritaceiventer 33 101.5 7.8 48.5 2.6 45.5 4.1 18.8 0.8 13 0.9 107 Hemitriccus minor 26 96.5 4.5 50.5 2.6 44.5 2.8 15.2 1 11.4 0.7 108 Hemitriccus mirandae 0 ------109 Hemitriccus nidipendulus 3 95.7 0.6 44 2 40.7 4 18.2 0.5 11.6 0.4 110 Hemitriccus obsoletus 2 115.5 2.1 56.5 0.7 54 1.4 20.3 0.8 13.5 0.8 111 Hemitriccus orbitatus 4 111 6.1 51.5 1.7 48.8 6.3 17.9 1 12.2 0.7 112 Hemitriccus rufigularis 14 109.9 6.9 51.6 3.2 52.1 4.9 16.6 1.2 12.8 1 113 Hemitriccus spodiops 5 90.2 3.6 46.4 3.2 38 6.5 15.1 0.5 10.7 1.2 114 Hemitriccus striaticollis 18 101.3 5.5 49.4 3 43.6 3.2 17.4 0.8 12.3 0.8 115 Hemitriccus zosterops 41 104.3 7.4 50.2 3.3 48.4 4.8 15.2 1 11.6 0.9 116 Heteroxolmis dominicana 8 211 6.3 111.1 4.8 100.9 4.3 26 0.9 25.1 1.4 117 Hirundinea bellicosa 4 173.8 8.5 108.3 4.4 78 4.3 13.3 1.2 14.6 1.2 118 Hirundinea ferruginea 10 175.3 6.4 113 4.8 84.7 4.5 14.1 0.8 15.7 1 119 Hymenops perspicillatus 13 156.9 8.2 89.4 3.2 65.5 4.1 27.5 1.2 20.5 1.2 120 Inezia inornata 11 100.5 5.1 47 2.3 47.2 3.3 15.2 0.4 11.3 0.8 170

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 91 Fluvicola albiventer * 10 18.2 1.1 8.3 1.1 4.9 0.2 3.4 0.3 92 Fluvicola nengeta * 7 19 0.9 8.1 0.9 5.2 0.4 3.7 0.4 93 Fluvicola pica * 10 18 1 9.3 0.9 5.2 0.4 4 0.4 Griseotyrannus 94 aurantioatrocristatus 9 17.7 1.2 12.8 0.8 7.4 0.5 5.2 0.3 95 Gubernetes yetapa * 4 28.2 0.4 17.3 1.1 10.6 0.4 8.6 0.3 96 Hemitriccus aenigma 6 14 0.6 7.6 0.8 4 0.1 3 0.2 97 Hemitriccus cinnamomeipectus 6 15 0.5 7.9 0.7 4 0.3 3.2 0.2 98 Hemitriccus diops 8 13.8 0.7 8 0.9 4 0.2 3.4 0.4 99 Hemitriccus flammulatus 31 14.5 0.6 8.5 0.7 4.6 0.2 3.4 0.3 100 Hemitriccus furcatus 1 12.5 - 6.4 - 3.8 - 2.9 - 101 Hemitriccus granadensis 23 13.4 0.4 6.7 0.6 3.7 0.3 3.1 0.2 102 Hemitriccus inornatus 0 ------103 Hemitriccus iohannis 5 14 0.4 8.6 1 4.4 0.3 3.6 0.2 104 Hemitriccus josephinae 0 ------105 Hemitriccus kaempferi 0 ------106 Hemitriccus margaritaceiventer 33 14.5 0.5 7.7 0.5 4.1 0.2 3.4 0.3 107 Hemitriccus minor 26 14.5 0.7 6.9 0.6 4.1 0.3 3.1 0.3 108 Hemitriccus mirandae 0 ------109 Hemitriccus nidipendulus 3 13.9 0.2 7.6 0.2 3.8 0.1 3 0.5 110 Hemitriccus obsoletus 2 14.1 0 8.2 0.1 3.5 - 3.7 0.6 111 Hemitriccus orbitatus 4 14.4 0.5 8.1 0.2 4.3 0.2 3.4 0.1 112 Hemitriccus rufigularis 14 14.7 0.7 8 0.6 4.1 0.2 3.2 0.3 113 Hemitriccus spodiops 5 13.1 0.3 6.6 0.8 3.8 0.1 3.4 0.2 114 Hemitriccus striaticollis 18 14.7 0.6 8.1 0.7 4.2 0.2 3.6 0.3 115 Hemitriccus zosterops 41 13.5 0.5 8.4 0.4 4.2 0.2 3.2 0.2 116 Heteroxolmis dominicana 8 22.7 1 13.3 0.8 6.6 0.2 5.7 0.4 117 Hirundinea bellicosa 4 20.6 2 11.9 0.3 8.6 0.5 4.7 0.9 118 Hirundinea ferruginea 10 22.7 1.4 13.4 1.2 10.3 0.6 5 0.5 119 Hymenops perspicillatus 13 19.2 1 10.2 0.6 4.6 0.2 4.6 0.4 120 Inezia inornata 11 12 0.6 6.2 0.7 3.1 0.1 2.6 0.3 171

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 121 Inezia subflava 10 108.9 2.8 49.9 4.1 50.6 3.5 15.7 0.8 12.5 0.9 122 Inezia tenuirostra 0 ------123 Knipolegus aterrimus * 8 162.4 6.5 83.9 2.4 75.6 2.9 23 2.1 16.9 0.7 124 Knipolegus cyanirostris * 10 151.9 5 79.4 3.6 72.7 3.7 19.7 1.2 15.4 1.3 125 Knipolegus hudsoni * 3 148.3 10.4 70 1 70.3 3.2 20 1 15.1 0.3 126 Knipolegus lophotes * 2 201.5 2.1 113 7.1 91 8.5 25 0.8 19.8 0.2 127 Knipolegus nigerrimus * 2 171.5 7.8 101.5 3.5 86 5.7 23.8 0.8 17 0.5 128 Knipolegus orenocensis * 3 154 7 79.7 5 74.7 1.2 20.1 0.5 16.4 0.9 129 Knipolegus poecilocercus * 10 141.5 6.4 73.4 8 64.9 5.3 17 0.8 14.7 0.7 130 Knipolegus poecilurus * 1 123 - 72 - 61 - 17.5 - 13.8 - 131 Knipolegus sigatus * 1 152 - 85 - 82 - 22.8 - 16.8 - 132 Knipolegus straiticeps * 10 127.9 3.5 56.9 0.7 59.6 1.3 19 0.8 14.6 0.7 133 Laniocera hypopyrra 1 203 - 107 - 90 - 19.8 - 19.3 - 134 Laniocera rufescens 1 204 - 114 - 96 - 21.6 - 17.1 - 135 Lathotriccus euleri 10 131.2 8.7 63.9 2.3 64.8 3.6 14.3 0.5 12.9 0.9 136 Lathotriccus griseipectus 2 124 0 57.5 0.7 59 2.8 14.9 1.6 12.9 0.1 137 Legatus leucophaius 10 146.8 8.8 84.9 4.1 65.6 3.5 15.6 0.8 16.1 0.9 138 Leptopogon amaurocephalus 11 122.8 5 63.3 2.5 59.7 6.3 14.9 0.9 11.3 0.6 139 Leptopogon rufipectus 10 128.2 6.7 67.2 4 63.9 3.8 16.2 0.7 13.3 0.9 140 Leptopogon superciliaris 10 127.3 6.4 63.6 4.2 60.1 5.3 14.5 0.9 12.6 0.7 141 Leptopogon taczanowski 10 122.8 5.3 64.4 2.5 60.1 3.5 14.6 0.4 12.7 0.3 142 Lessonia oreas 6 125.5 3.3 79 2.6 50.7 3.9 22.4 1 16.2 0.9 143 Lessonia rufa 9 122.2 7.2 71.3 3.5 45.2 3.5 21.1 1.2 16.4 0.9 144 Lophotriccus eulophotes 10 93.5 5.7 47 1.2 42.9 3.6 14.5 0.7 11.2 0.4 145 Lophotriccus galeatus 10 94.2 6.2 45.4 1.9 42.1 4.3 15.3 0.6 10.5 0.9 146 Lophotriccus pileatus 10 93.7 5.9 48.8 3.3 38.3 5.5 15.1 0.7 11 0.6 147 Lophotriccus vitiosus 10 90.4 6.9 46.6 2.5 39.1 4.6 14 0.6 11.2 0.8 148 Machetornis rixosus * 10 186.6 9.9 91.8 4.8 84.5 4.7 29.2 2.1 23.1 3.4 149 Mecocerculus calopterus * 8 107.1 4.6 53.8 3.1 50.6 2.9 16.8 0.9 11.7 0.7 150 Mecocerculus hellmayri * 8 98.3 3.9 56.5 4.2 44.4 2.7 15.8 0.7 12.7 0.4

172

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 121 Inezia subflava 10 13.5 0.7 7.9 0.9 4.2 0.5 3.3 0.4 122 Inezia tenuirostra 0 ------123 Knipolegus aterrimus * 8 19.6 1.3 10.3 0.7 5.1 0.1 4.4 0.4 124 Knipolegus cyanirostris * 10 16.9 0.7 9.7 0.7 5.5 0.2 4.5 0.3 125 Knipolegus hudsoni * 3 17.1 0.4 10 0.4 5.2 0.1 4.4 0.2 126 Knipolegus lophotes * 2 22.8 0.4 11.8 1.8 5.8 - 5.5 0.1 127 Knipolegus nigerrimus * 2 18 1.2 10.5 1.6 5.9 - 5 0.2 128 Knipolegus orenocensis * 3 18.9 1.8 11.1 0.9 5.9 0.4 4.8 0.2 129 Knipolegus poecilocercus * 10 16 0.7 9.6 0.4 5.2 0.2 4.1 0.2 130 Knipolegus poecilurus * 1 13 - 7.8 - - - 3.3 - 131 Knipolegus sigatus * 1 17.2 - 8.9 - - - 4.3 - 132 Knipolegus straiticeps * 10 14.8 1 9.2 1 4.4 0.2 3.7 0.3 133 Laniocera hypopyrra 1 19.3 - 16 - 6.1 - 6.1 - 134 Laniocera rufescens 1 20.8 - 15.3 - 6.6 - 6.6 - 135 Lathotriccus euleri 10 14.9 0.9 9.1 0.4 5.8 0.5 4 0.5 136 Lathotriccus griseipectus 2 14 0.4 8.1 0.4 5.6 - 3.6 0.2 137 Legatus leucophaius 10 14.8 0.6 12.4 0.8 7 0.6 5 0.4 138 Leptopogon amaurocephalus 11 14.8 0.4 8.5 0.8 4 0.2 4.3 0.3 139 Leptopogon rufipectus 10 13.8 0.7 9 0.5 3.7 0.5 4 0.2 140 Leptopogon superciliaris 10 15.1 0.8 9.2 0.6 4.3 0.4 4.1 0.2 141 Leptopogon taczanowski 10 12.7 0.3 8.3 0.6 3.4 0.3 4 0.3 142 Lessonia oreas 6 14.5 0.7 7 0.6 3.5 0.3 3.2 0.1 143 Lessonia rufa 9 14.1 0.7 7.2 1.1 3.5 0.4 3.2 0.3 144 Lophotriccus eulophotes 10 12.3 0.5 7.6 0.6 3.7 0.2 3.2 0.3 145 Lophotriccus galeatus 10 12.1 0.4 7.2 0.4 3.9 0.2 3.1 0.3 146 Lophotriccus pileatus 10 12.7 0.5 7.2 0.4 3.5 0.2 3.1 0.3 147 Lophotriccus vitiosus 10 12.3 0.6 7.6 0.6 3.8 0.3 3.3 0.4 148 Machetornis rixosus * 10 23.7 1.8 11.8 0.9 6.1 0.6 5.1 0.3 149 Mecocerculus calopterus * 8 12.7 0.5 6.3 0.5 3.1 0.2 3.2 0.3 150 Mecocerculus hellmayri * 8 11.3 0.3 6.3 0.5 2.8 0.3 3.2 0.3

173

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 151 Mecocerculus leucophrys * 9 142.1 3.3 68.6 3.8 75.3 3.3 22.1 1.3 14.6 0.9 152 Mecocerculus minor * 11 116.8 5.4 57.5 3.1 58.9 4.9 16.8 1.1 11.7 0.9 153 Mecocerculus poecilocercus * 14 102.5 3.5 55.9 2.1 49.2 2.8 15.9 0.7 12.4 0.9 154 Mecocerculus stictopterus * 10 115.1 3.2 61.1 1.9 58.9 2.2 17.4 0.7 13 0.9 155 Megaynchus pitangua * 10 217.7 7.5 122 3.3 96.1 4.6 20.5 0.9 24.3 0.9 156 Mionectes macconnelli 11 118.5 4.9 64.7 5.1 54.4 3.6 15.3 0.6 12.8 0.9 157 Mionectes oleagineus 32 116.5 8.2 59.9 3.7 49.5 5 15.1 1.2 11.7 1 158 Mionectes olivaceus 16 127.6 7.3 63.9 3.2 52.3 2.3 16.6 0.7 12.6 1.2 159 Mionectes rufiventris 11 130.8 9.2 66.8 2.8 60.9 3.7 16.6 0.6 13.7 0.6 160 Mionectes striaticollis 10 122.4 8.3 66.3 3.9 55.5 4.2 16.6 0.6 13.9 0.7 161 Mitrephanes olivaceus 1 120 - 68 - 61 - 12.2 - 11 - 162 Mitrephanes phaeocercus 14 124.2 4.2 68.1 4.1 64.1 3.1 12.7 1 10.5 0.5 163 Muscigralla brevicauda * 7 106.6 6 59.4 2.9 36.1 1.8 25.3 1.1 17.4 1 164 Muscipipra vetula 3 211 11.4 118 5.6 122 8 19.2 0.4 19.3 1.9 165 Muscisaxicola albifrons 3 211.3 11.6 152.7 6.5 96.7 3.8 35.8 2.7 21.2 6.6 166 Muscisaxicola albilora 8 169.5 10.2 112.4 5.5 77.1 4.8 30.7 1.6 19.8 1.1 167 Muscisaxicola alpina 3 175.3 8.5 119.3 7.6 89 6.1 31.5 1.1 20 0.7 168 Muscisaxicola capistrata 10 170 7.3 115.6 3.7 75.6 4.7 28.3 1.1 19 0.7 169 Muscisaxicola cinerea 3 172 8.5 106.3 3.5 76.7 3.1 28.9 0.6 18.4 2.5 170 Muscisaxicola flavinucha 10 171.9 4.8 122.5 2.8 75.2 2.4 29.9 1 19.9 1.5 171 Muscisaxicola fluviatilis 2 128.5 3.5 73 1.4 57.5 0.7 20.7 1.3 16 0.4 172 Muscisaxicola frontalis 4 179.8 3.1 123 7.7 81.8 5.7 32.1 0.4 21.1 1 173 Muscisaxicola juninensis 2 169 7.1 111 4.2 76.5 3.5 29.7 1.4 19.3 0.1 174 Muscisaxicola macloviana 10 158.9 8.1 100 3.1 68.1 3.2 28.1 1.4 19.1 1.1 175 Muscisaxicola maculirostris 10 143.2 9.3 84.1 3.3 66 2.7 25.7 1.5 16.6 0.6 176 Muscisaxicola rufivertex 10 172 8.6 111 7.4 79.4 5.1 29.5 1.5 18.1 0.8 177 Myiarchus antillarum 10 172.2 5.1 85.6 3.6 83.5 4.2 21.9 0.4 16.7 0.9 178 Myiarchus apicalis * 10 187.3 5.3 91.1 2.5 91.9 2.8 21.1 0.5 17 1.1 179 Myiarchus barbirostris * 10 148.2 6.8 71.8 3.3 73.8 2.7 18 0.8 13.8 0.9 180 Myiarchus cephalotes 10 189.2 4.3 89.9 3 93.2 2.7 20.3 1.3 15.7 1.2

174

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 151 Mecocerculus leucophrys * 9 14.4 0.8 8 1.1 3.7 0.9 3.5 0.2 152 Mecocerculus minor * 11 12.2 0.4 6.2 0.5 3 0.4 3.4 0.2 153 Mecocerculus poecilocercus * 14 11.2 0.5 6.1 0.7 2.6 0.3 3.1 0.3 154 Mecocerculus stictopterus * 10 12.3 0.7 6.6 0.5 2.9 0.3 3.1 0.3 155 Megaynchus pitangua * 10 36.9 1.8 17.9 1 13.7 0.8 10.7 0.4 156 Mionectes macconnelli 11 13.9 0.8 8.8 0.5 4 0.2 3.6 0.3 157 Mionectes oleagineus 32 13.4 1.2 8.6 0.8 3.7 0.3 3.6 0.2 158 Mionectes olivaceus 16 15.2 0.4 8.9 0.6 4 0.3 3.7 0.3 159 Mionectes rufiventris 11 14.7 0.7 9.2 0.6 4.2 0.3 3.8 0.1 160 Mionectes striaticollis 10 14.3 0.6 8.5 0.7 3.8 0.2 3.8 0.2 161 Mitrephanes olivaceus 1 13 - 7 - 3 - - - 162 Mitrephanes phaeocercus 14 13.2 0.5 6.7 0.7 4.8 0.4 3 0.3 163 Muscigralla brevicauda * 7 16.5 1.1 8.9 0.8 4.3 0.5 3.7 0.3 164 Muscipipra vetula 3 19.4 1 11.9 1.2 7.6 0.8 5 0.3 165 Muscisaxicola albifrons 3 25.6 0.9 11.6 0.8 4.9 0.3 5.3 0.9 166 Muscisaxicola albilora 8 21 1.5 9.8 0.9 4.2 0.1 4 0.2 167 Muscisaxicola alpina 3 19.4 1.5 8.9 0.8 4.3 0 4.1 0.1 168 Muscisaxicola capistrata 10 20 1.2 9.2 0.7 4.4 0.2 4 0.3 169 Muscisaxicola cinerea 3 13.5 5.1 8.8 0.4 3.9 0.4 3.3 0.8 170 Muscisaxicola flavinucha 10 18.6 1.5 10.4 0.8 4.3 0.4 4.1 0.3 171 Muscisaxicola fluviatilis 2 16.1 0.3 7.3 0.9 4.4 - 4 0.8 172 Muscisaxicola frontalis 4 23.2 0.9 10 0.9 4.1 0.1 4.2 0.3 173 Muscisaxicola juninensis 2 19.4 0.8 9.7 0.9 4.3 - 3.6 0.1 174 Muscisaxicola macloviana 10 17.5 0.5 8.4 0.8 3.6 0.4 3.6 0.3 175 Muscisaxicola maculirostris 10 17.3 0.8 8.8 0.7 3.7 0.4 3.5 0.2 176 Muscisaxicola rufivertex 10 21 1.3 9 0.7 4 0.2 3.9 0.5 177 Myiarchus antillarum 10 23.3 1 12.3 0.5 7.1 0.3 5.6 0.4 178 Myiarchus apicalis * 10 23 1.3 12 0.6 8 0.3 5.8 0.4 179 Myiarchus barbirostris * 10 19.9 0.8 10.1 0.9 7.1 0.4 4.5 0.4 180 Myiarchus cephalotes 10 21.9 0.6 11.9 0.7 7.5 0.3 5.6 0.2

175

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 181 Myiarchus cinerascens 12 192.5 8 96.5 4.3 98.1 4.5 22.4 0.8 17.3 1.1 182 Myiarchus crinitis 10 188.6 4.2 97.2 3.6 93.1 4.6 20.7 0.7 18.2 0.8 183 Myiarchus ferox 10 180.6 8.1 86.5 1.6 90.3 4.9 21.2 0.6 17 1.2 184 Myiarchus magnirostris 10 145.2 5.3 68.3 2.4 65.3 3.1 20.7 0.3 14.5 1 185 Myiarchus nugator 10 207.6 5.6 98.9 3.8 95.8 4.6 24.4 0.7 18.8 1.1 186 Myiarchus nuttingi 10 176.9 8.4 90.8 5.3 90.4 4.6 21.3 1.5 16.7 0.9 187 Myiarchus oberi 10 188.5 5.7 98 4.9 90.3 3.6 23.5 0.7 18.5 0.8 188 Myiarchus panamensis 10 186.6 6 91.5 4.6 93.4 4.1 22.2 1.8 16.6 0.8 189 Myiarchus phaeocephalus 8 187.5 4.9 92.8 4.3 92.3 5.5 22.3 1.4 16.7 1.2 190 Myiarchus sagrae 10 171.5 5.7 81.3 3.9 81.9 1.7 19.6 0.7 16 1.1 191 Myiarchus semirufus * 1 173 - 84 - 88 - 20 - 16.9 - 192 Myiarchus stolidus 10 173.9 6 81.1 2.6 83.7 3.8 20.4 0.7 16.8 1.7 193 Myiarchus swainsoni * 4 178.3 21.5 90.3 5.9 86 4.9 19.5 1 16.7 0.6 194 Myiarchus tuberculifer * 10 159.2 9.5 78.4 3.7 75.4 5.9 19 0.7 15.5 0.4 195 Myiarchus tyrannulus 10 185.7 10 96.6 6.2 92.7 3.2 21.9 1.3 18.2 1.3 196 Myiarchus validus 9 209.2 10.6 100.2 3.5 102.7 4.7 23.4 1.4 19.7 1.1 197 Myiarchus venezuelensis * 10 182.7 4.5 87.4 2 88.8 2.9 21.1 0.7 16 0.9 198 Myiarchus yucatanensis * 10 170.2 6.6 81.8 3.6 84.5 5.6 21.6 0.8 16.4 0.4 199 Myiobius atricaudus 10 124.9 3.3 59.3 1.8 63.5 1.4 18.2 1.3 12.8 0.6 200 Myiobius barbatus 14 121.1 5.4 63.1 2.8 58.8 3.7 16.2 1.1 12.8 0.6 201 Myiobius erythrurus 10 96.8 3.5 49.4 2 41.9 2.9 14.7 1.1 12.2 1.2 202 Myiobius villosus 6 135.7 3.1 69.8 1.8 64.7 2.3 18.4 0.8 13.8 0.8 203 Myiodynastes bairdii * 3 212 5.6 114.3 2.9 104.7 3.2 24.2 0.4 23.1 1.5 204 Myiodynastes chrysocephalus * 4 203 3.7 109 3.9 99.8 3.8 18 0.7 18.6 0.8 205 Myiodynastes hemichrysus * 6 199 4.6 103.2 2.2 94 1.7 18.5 0.9 18.5 0.7 206 Myiodynastes luteiventris * 20 190 8.6 113.3 3.6 88.3 3.9 19.1 1.1 19.6 1.5 207 Myiodynastes maculatus * 10 202.9 7.5 104.2 5.1 90.7 2.4 19.5 0.5 19.7 0.5 208 Myiopagis caniceps * 10 122.6 7 60.8 1.7 58.5 3.1 16.7 0.6 12.7 1.9 209 Myiopagis cotta * 6 134.5 5.5 65.8 2.9 64.8 2.7 16.8 0.7 13.1 1 210 Myiopagis flavivertex * 1 122 - 53 - 50 - 15.2 - 12.3 -

176

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 181 Myiarchus cinerascens 12 24.1 1.2 11.6 0.5 7.3 0.2 6.4 0.4 182 Myiarchus crinitis 10 25.4 1.3 13.2 0.6 8.5 0.3 7.1 0.5 183 Myiarchus ferox 10 23.5 0.8 12.1 0.5 7.9 0.5 5.8 0.3 184 Myiarchus magnirostris 10 19.1 0.8 10.2 0.7 6.7 0.3 4.4 0.2 185 Myiarchus nugator 10 26.8 1.1 15.4 1.3 8.9 0.7 6.7 0.4 186 Myiarchus nuttingi 10 22.4 1.4 11.6 0.7 6.9 0.3 5.9 0.4 187 Myiarchus oberi 10 26.6 0.9 13.1 1.1 7.8 0.6 5.9 0.3 188 Myiarchus panamensis 10 23.9 0.9 13.3 1 8.2 0.4 6.1 0.4 189 Myiarchus phaeocephalus 8 23.8 1.4 12.6 0.9 7.6 0.6 5.6 0.2 190 Myiarchus sagrae 10 22.8 0.9 12.4 3.2 7.5 0.3 5.4 0.5 191 Myiarchus semirufus * 1 25.5 - 13 - 7 - 5.3 - 192 Myiarchus stolidus 10 22.6 1.2 10.9 1.1 7.4 0.6 5.2 0.2 193 Myiarchus swainsoni * 4 22.6 0.5 10.9 0.2 7.4 0.4 5.7 0.3 194 Myiarchus tuberculifer * 10 21.4 1.1 10.7 1.1 7.1 0.4 5.2 0.4 195 Myiarchus tyrannulus 10 25 1.8 13.2 1 8.5 0.6 6.6 0.7 196 Myiarchus validus 9 26.4 1.2 15.4 0.6 9 0.4 7 0.4 197 Myiarchus venezuelensis * 10 22.7 0.9 12.6 0.6 7.6 0.4 5.9 0.3 198 Myiarchus yucatanensis * 10 21.5 0.8 12.4 0.8 7.5 0.3 5.9 0.4 199 Myiobius atricaudus 10 13.9 0.6 7.9 0.6 4.7 0.7 3.2 0.3 200 Myiobius barbatus 14 14.1 0.5 8.7 0.4 4.9 0.3 3.4 0.2 201 Myiobius erythrurus 10 10.5 0.6 7.7 0.5 4.1 0.5 2.9 0.3 202 Myiobius villosus 6 16.4 0.3 9.6 0.2 5.6 0.2 3.6 0.1 203 Myiodynastes bairdii * 3 30.2 0.7 18.1 1.6 10.6 0.6 8.9 0.5 204 Myiodynastes chrysocephalus * 4 27.2 1.1 16.9 1 11.4 0.4 8.4 0.7 205 Myiodynastes hemichrysus * 6 25.2 1.4 16.4 1.3 11.1 0.4 7.4 0.4 206 Myiodynastes luteiventris * 20 25.9 0.9 16.6 0.8 10.2 0.6 8.4 0.6 207 Myiodynastes maculatus * 10 28.2 1.6 17.1 1 11.5 0.5 8.7 0.4 208 Myiopagis caniceps * 10 12.1 1 7.6 1 3.9 0.3 3.7 0.2 209 Myiopagis cotta * 6 13.3 1.2 7.8 0.7 3.7 0.3 3.1 0.4 210 Myiopagis flavivertex * 1 13 - 8.8 - - - 3.1 -

177

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 211 Myiopagis gaimardii * 11 115.9 6.9 55.7 3 56.5 4.7 16.9 1.3 12.8 1 212 Myiopagis subplacens * 10 143.8 8.7 68.6 3.8 69.8 5 20 1.4 13.1 1.7 213 Myiopagis viridicata * 12 133.9 7.7 64.1 2.6 65.5 4 17.1 1.1 12.5 1.1 214 Myiophobus cryptoxanthus 1 121 - 59 - 57 - 15.9 - 13.8 - 215 Myiophobus fasciatus 10 128.2 10.3 61.6 1.9 59.9 4.9 15.9 1 13.5 0.6 216 Myiophobus flavicans 10 126.9 6.1 69.1 3.1 58.1 1.8 17.4 0.6 14.3 0.6 217 Myiophobus inornatus 1 103 - 55 - 47 - 15.9 - 13 - 218 Myiophobus lintoni 3 117 2.6 59.7 2.3 58 4.4 15.7 0.6 12.5 0.4 219 Myiophobus ochraceiventris 9 129.8 5.6 66.2 3.6 70 4.1 16.8 1.2 14 0.7 220 Myiophobus phoenicomitra 6 120.2 4.6 62 4 59.3 3.7 16.2 0.9 12.4 0.4 221 Myiophobus pulcher 4 109.5 8.6 56.5 5.8 49.3 7.1 15.7 0.7 12.7 1.2 222 Myiophobus roraimae 11 131.9 7.2 68.8 2.8 66.1 3.3 17.8 0.6 13.7 0.4 223 Myiornis albiventris 9 77.6 4 37.1 2 26.9 1.8 13.3 0.6 9.6 0.6 224 Myiornis atricapillus 10 64.6 3.1 35.4 2.6 16.3 2.2 12.7 0.6 10.3 0.5 225 Myiornis auricualris 2 80 0 37.5 2.1 33 14.1 13.9 1.6 9.9 1 226 Myiornis ecaudatus 12 62 3.5 31.8 1.6 16.2 2.4 11.5 0.8 9.6 0.7 227 Myiotheretes fumigatus 10 198.2 8.3 106.9 3.3 90.8 3.9 23.5 1.1 20.4 0.8 228 Myiotheretes fuscorufus 1 173 - 103 - 84 - 20.5 - 17.7 - 229 Myiotheretes pernix 10 191.9 3.8 99.7 4.3 87.4 3.6 23.1 0.8 21.2 0.8 230 Myiotheretes striaticollis 10 225.8 5.8 137.5 5.7 102.8 4 24.8 1.2 23.2 0.8 231 Myiotriccus ornatus 10 118.8 9.3 65.8 4 54.2 6.7 15.8 1.2 13.1 0.9 232 Myiozetetes cayanensis * 10 165.2 10.3 85.9 4.7 74.7 3.1 18.5 0.7 17.6 1.2 233 Myiozetetes granadensis * 10 165.6 6.4 85.2 2.8 76.1 2.7 17.6 0.7 17.6 1.1 234 Myiozetetes luteiventris * 5 138.8 7.6 76.4 10.5 67.2 3 14.6 0.6 14 0.7 235 Myiozetetes similis * 10 172.4 8.2 91.5 4.8 81.5 3.2 19.3 1.3 17.5 1 236 Neoxolmis rufiventris 7 235.9 7.7 159.7 14.2 105.6 4.9 35.3 4.1 27.7 1.6 237 Nesotriccus ridgwayi 5 127 7.6 57.6 3.7 56.2 1.9 19.9 1.1 13.9 0.9 238 Ochthoeca cinnamomeiventris * 10 124.2 6.5 66.5 2.2 56 2.1 18.1 0.7 14.3 0.7 239 Ochthoeca fumicolor * 10 152.1 5.7 82.5 2.6 74.6 2.3 22.5 0.4 16.7 1.7 240 Ochthoeca leucophrys * 10 151.1 5.3 76.8 3.4 76.2 3 21.6 0.6 15.9 1

178

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 211 Myiopagis gaimardii * 11 12.7 0.7 8.2 0.9 - - 3.5 0.3 212 Myiopagis subplacens * 10 14.6 0.7 8.2 0.9 - - 4 0.2 213 Myiopagis viridicata * 12 13.2 0.8 7.9 0.9 - - 3.4 0.2 214 Myiophobus cryptoxanthus 1 13.3 - 8.6 - - - 3.4 - 215 Myiophobus fasciatus 10 14.3 0.6 8.3 0.6 5.5 0.2 3.8 0.3 216 Myiophobus flavicans 10 13.6 0.7 8.2 0.5 5.1 0.1 3.9 0.3 217 Myiophobus inornatus 1 12.8 - 8 - - - 3.2 - 218 Myiophobus lintoni 3 12.5 0.1 7.4 0.6 4 0.2 3.2 0.3 219 Myiophobus ochraceiventris 9 13.2 0.4 8.1 0.4 4.4 0.2 3.4 0.3 220 Myiophobus phoenicomitra 6 13.3 0.4 8.1 0.7 4.8 0.1 3.8 0.2 221 Myiophobus pulcher 4 11.4 0.5 6.2 0.9 4.5 0.4 3.3 0.4 222 Myiophobus roraimae 11 14.4 0.7 8.9 0.6 5 0.3 3.9 0.3 223 Myiornis albiventris 9 11.5 0.7 6.3 0.7 3.5 0.3 2.7 0.2 224 Myiornis atricapillus 10 11.7 0.6 6.7 0.6 3.7 0.3 2.8 0.3 225 Myiornis auricualris 2 11.9 0.1 6.5 0.1 3.6 - 3 0.3 226 Myiornis ecaudatus 12 11.5 0.5 6.2 0.4 3.5 0.2 2.8 0.2 227 Myiotheretes fumigatus 10 24.2 0.9 13.1 1 7.6 0.5 5.9 0.3 228 Myiotheretes fuscorufus 1 22.7 - 12 - - - 5.4 - 229 Myiotheretes pernix 10 25.9 0.9 13.3 0.7 7.7 0.5 6 0.2 230 Myiotheretes striaticollis 10 29.1 1.1 15.6 0.7 9 0.6 7.9 0.3 231 Myiotriccus ornatus 10 14.2 0.7 8.8 0.8 5.1 0.5 4.1 0.3 232 Myiozetetes cayanensis * 10 17 0.9 11 0.4 5.7 0.3 5.2 0.2 233 Myiozetetes granadensis * 10 17.9 1.3 11.9 0.8 6.4 0.5 5.1 0.4 234 Myiozetetes luteiventris * 5 13.5 0.9 9.8 0.7 5.3 0.6 4.3 0.3 235 Myiozetetes similis * 10 17.4 1 11.5 0.9 5.9 0.4 5.2 0.4 236 Neoxolmis rufiventris 7 23.9 1.4 13.5 2 6.9 0.3 5.9 0.5 237 Nesotriccus ridgwayi 5 18 0.7 8.4 0.6 4.7 0.4 3.6 0.3 238 Ochthoeca cinnamomeiventris * 10 14.5 0.7 8.3 0.8 5.1 0.4 3.8 0.3 239 Ochthoeca fumicolor * 10 16.2 0.7 9.1 0.8 4.7 0.3 3.9 0.2 240 Ochthoeca leucophrys * 10 17 0.7 8.8 0.8 4.4 0.3 3.9 0.3

179

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 241 Ochthoeca oenanthoides * 5 158.8 7.9 85.4 3.5 75 3.7 21.7 0.7 16.1 1.5 242 Ochthoeca piurae * 1 104 - 61 - 43 - 18.2 - 13.7 - 243 Ochthoeca rufipectoralis * 10 133.3 5.4 71.5 3.6 67 3.2 18 1 14.3 0.8 244 Ochthoeca salvini * 2 134 2.8 73.5 3.5 64 2.8 17.4 1.6 14.7 1.3 245 Ochthornis littoralis 3 127.3 1.5 65 7.2 63.3 4 17.4 1.1 15.4 0.9 246 Oncostoma cinereigulare 10 91.5 7.4 47.9 4 41.2 4.5 14.6 0.8 10 0.8 247 Oncostoma olivaceum 10 90.2 7.3 45.3 2.7 37.9 3.5 14.7 0.7 10.4 1 248 Onychorhynchus coronatus 10 162.8 16.7 84.7 7.5 73.5 6.9 16.5 0.8 15.5 0.8 249 Ornithion brunneicapillum 10 81.3 3.1 44.8 1.8 28.2 1.4 13.4 0.6 11.2 0.6 250 Ornithion inerme 9 86.7 4.9 44.7 2.6 34.2 3.5 13.1 0.5 10.7 0.6 251 Ornithion semiflavum 10 79.2 2.4 46.1 1.3 28.7 3.9 13.8 0.6 11.6 0.4 252 Pachyramphus aglaiae 1 180 - 92 - 76 - 22.4 - 18 - 253 Pachyramphus albogriseus 1 137 - 81 - 60 - 18.7 - 15.2 - 254 Pachyramphus castaneus 1 136 - 74 - 58 - 18.4 - 15.9 - 255 Pachyramphus cinnamomeus 1 140 - 77 - 67 - 21.4 - 15.7 - 256 Pachyramphus homochrous 1 154 - 90 - 66 - 22.3 - 19.8 - 257 Pachyramphus major 1 144 - 78 - 67 - 19.5 - 17.5 - 258 Pachyramphus marginatus 1 125 - 68 - 55 - 16.9 - 14.8 - 259 Pachyramphus minor 1 163 - 92 - 71 - 20 - 18.7 - 260 Pachyramphus niger 1 185 - 96 - 75 - 23.4 - 16 - 261 Pachyramphus polychopterus 1 136 - 74 - 61 - 18.8 - 14.4 - 262 Pachyramphus rufus 1 126 - 68 - 52 - 17.6 - 14.5 - 263 Pachyramphus spodiurus 1 143 - 72 - 59 - 20.3 - 15.1 - 264 Pachyramphus surinamus 0 ------265 Pachyramphus validus 1 172 - 105 - 75 - 22.2 - 18.5 - 266 Pachyramphus versicolor 1 115 - 65 - 49 - 15.8 - 14.2 - 267 Pachyramphus viridis 1 141 - 76 - 63 - 17.8 - 15.9 - 268 Phaeomyias murina * 11 117.6 5.8 60.8 3 59.1 4.5 19.8 1.2 13.3 0.6 269 Phelpsia inornata 3 179 6.6 94.7 5 83 5.3 22.6 1.6 18.7 1.5 270 Philohydor lictor 1 163 - 84 - 77 - 27.9 - 15.2 -

180

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 241 Ochthoeca oenanthoides * 5 17.6 0.5 9.9 0.5 4.6 0.4 3.8 0.3 242 Ochthoeca piurae * 1 14.8 - 8 - 3.7 - 3.4 - 243 Ochthoeca rufipectoralis * 10 13.7 0.8 7.1 0.7 4.2 0.3 3.4 0.5 244 Ochthoeca salvini * 2 16.7 0.2 9 0.9 5.1 0.8 3.9 0.5 245 Ochthornis littoralis 3 15.7 0.3 9.2 0.8 5.2 0.5 3.8 0.1 246 Oncostoma cinereigulare 10 13.1 0.4 7 0.5 3.8 0.2 3.3 0.2 247 Oncostoma olivaceum 10 13.6 0.3 7.8 0.7 4.2 0.2 3.7 0.3 248 Onychorhynchus coronatus 10 26.5 3.1 10.9 0.8 8.7 0.8 4.9 0.4 249 Ornithion brunneicapillum 10 10.2 0.4 6.7 0.5 3.7 0.3 3.4 0.2 250 Ornithion inerme 9 10.6 0.5 6.4 0.6 3.1 0.4 3.1 0.2 251 Ornithion semiflavum 10 9.8 0.3 6.9 0.5 3.3 0.3 3.5 0.2 252 Pachyramphus aglaiae 1 20.6 - 16.3 - 7.9 - 7.3 - 253 Pachyramphus albogriseus 1 12.2 - 13.3 - 6.7 - 5.8 - 254 Pachyramphus castaneus 1 17.1 - 9.5 - 7 - 4.6 - 255 Pachyramphus cinnamomeus 1 16.2 - 10.6 - 6.9 - 5.1 - 256 Pachyramphus homochrous 1 20.8 - 15.2 - 7.6 - 7.3 - 257 Pachyramphus major 1 16.1 - 14.1 - 6.5 - 5.4 - 258 Pachyramphus marginatus 1 14.3 - 10.7 - 6.8 - 5 - 259 Pachyramphus minor 1 19 - 15.5 - 6.8 - 7.8 - 260 Pachyramphus niger 1 22 - 16.2 - 8 - 7.1 - 261 Pachyramphus polychopterus 1 16.5 - 12 - 7 - 5.6 - 262 Pachyramphus rufus 1 15.4 - 9.5 - 6.3 - 5.3 - 263 Pachyramphus spodiurus 1 16.9 - 11.2 - 6.5 - 5.2 - 264 Pachyramphus surinamus 0 ------265 Pachyramphus validus 1 21 - 16.7 - 9.4 - 7.9 - 266 Pachyramphus versicolor 1 13.7 - 9 - 4.2 - 4.1 - 267 Pachyramphus viridis 1 15.2 - 9.3 - 5.8 - 4.9 - 268 Phaeomyias murina * 11 12.7 0.8 7.8 0.5 3.7 0.1 3.4 0.4 269 Phelpsia inornata 3 18.6 2.2 12.9 1.8 6.4 0.4 5.2 0.5 270 Philohydor lictor 1 22.9 - 10.3 - 4.5 - 5.1 -

181

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 271 Phyllomyias burmeisteri 4 116.5 8.4 62 2.9 50 5.7 13 1.2 13.4 0.6 272 Phyllomyias cinereiceps 10 102.5 4.1 61 2.4 45.2 1.9 13.8 0.9 12.6 1 273 Phyllomyias fasciatus 4 110.8 9.3 55 2 49.8 2.1 14.1 0.7 11.8 0.5 274 Phyllomyias griseiceps 10 101.5 6.5 51.6 2.5 47.2 3.8 14.1 0.8 10.8 1 275 Phyllomyias griseocapilla 3 111.7 3.8 56.3 4.5 52.3 0.6 15.9 1.7 12.9 1.5 276 Phyllomyias nigrocapillus 10 107.6 2.9 62.3 2.2 52.9 2.5 16.1 0.8 13.4 0.8 277 Phyllomyias plumbeiceps 6 110.8 5.6 58.5 3 55.5 3.6 15.4 1.3 11.4 0.5 278 Phyllomyias reiseri 0 ------279 Phyllomyias sclateri 4 122.5 5.4 60.5 2.4 60 3.2 16.2 0.7 12.5 1.1 280 Phyllomyias uropygialis 10 114.6 8 60.6 2.6 52.4 4.1 16.9 0.7 14.2 0.5 281 Phyllomyias virescens 10 127.2 2.8 58.8 3.7 58.4 4 16.3 0.9 12.8 0.7 282 Phyllomyias zeledoni 3 101.3 1.2 58 2.6 44.7 2.1 14.6 1.2 11.5 1 283 Phylloscartes ceciliae 0 ------284 Phylloscartes chapmani 0 ------285 Phylloscartes difficilis 1 110 - 52 - 59 - 18.8 - 13 - 286 Phylloscartes eximius 8 112.6 6 52.5 2.4 51.8 5.5 15.1 1.5 11.3 1.1 287 Phylloscartes flaviven 0 ------288 Phylloscartes flavovir 0 ------289 Phylloscartes gualaqui 0 ------290 Phylloscartes lanyoni 0 ------291 Phylloscartes nigrifro 0 ------292 Phylloscartes ophthalmicus 11 112.3 5.7 55.7 3.2 53 2.6 14.4 0.8 11.6 1 293 Phylloscartes orbitalis 10 104.8 5 52.9 2.4 49.7 3.2 15.3 0.8 11.6 0.6 294 Phylloscartes oustaleti 1 120 - 51 - 57 - 17.4 - 12.2 - 295 Phylloscartes paulist. 0 ------296 Phylloscartes poecilotis 10 109.8 7 56.9 2.9 54.8 4.8 15.6 0.9 11.6 0.8 297 Phylloscartes roquette 0 ------298 Phylloscartes superciliaris 0 ------299 Phylloscartes sylviolus 0 ------300 Phylloscartes venezuelanus 1 113 - 52 - 52 - 15.4 - 11.9 -

182

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 271 Phyllomyias burmeisteri 4 11.6 0.8 8.5 0.5 3.9 0.2 3.5 0.3 272 Phyllomyias cinereiceps 10 11 0.5 7 0.7 3.4 0.3 3.3 0.2 273 Phyllomyias fasciatus 4 11.2 0.4 6.3 0.9 - - 3 0.1 274 Phyllomyias griseiceps 10 10.3 0.4 7.2 0.8 4.4 0.4 3.1 0.3 275 Phyllomyias griseocapilla 3 10.7 0.4 6.6 1 3.9 0.7 3.4 0.3 276 Phyllomyias nigrocapillus 10 10.6 0.3 6.5 0.8 3.2 0.4 3.2 0.1 277 Phyllomyias plumbeiceps 6 11 0.4 6.6 0.8 3.5 0.8 3.2 0.3 278 Phyllomyias reiseri 0 ------279 Phyllomyias sclateri 4 11.4 0.3 6.9 0.5 3.4 0.3 3.1 0.2 280 Phyllomyias uropygialis 10 10.7 0.8 6.6 0.8 2.9 0.2 3.1 0.2 281 Phyllomyias virescens 10 11.7 0.7 7 0.8 3.5 0.5 3.5 0.3 282 Phyllomyias zeledoni 3 11.2 0.5 8 0.8 3.4 0.4 3.4 0.2 283 Phylloscartes ceciliae 0 ------284 Phylloscartes chapmani 0 ------285 Phylloscartes difficilis 1 12.7 - 6.7 - - - 3.2 - 286 Phylloscartes eximius 8 11 0.4 6.7 0.5 3.2 0.3 3.3 0.4 287 Phylloscartes flaviven 0 ------288 Phylloscartes flavovir 0 ------289 Phylloscartes gualaqui 0 ------290 Phylloscartes lanyoni 0 ------291 Phylloscartes nigrifro 0 ------292 Phylloscartes ophthalmicus 11 11.6 0.5 6.9 0.7 3.5 0.2 3 0.2 293 Phylloscartes orbitalis 10 12.5 0.6 6.4 0.7 3.6 0.2 3.2 0.2 294 Phylloscartes oustaleti 1 14.1 - 8.4 - - - 3.6 - 295 Phylloscartes paulist. 0 ------296 Phylloscartes poecilotis 10 12.1 0.4 6.4 0.5 3.4 0.3 3.2 0.3 297 Phylloscartes roquette 0 ------298 Phylloscartes superciliaris 0 ------299 Phylloscartes sylviolus 0 ------300 Phylloscartes venezuelanus 1 12.6 - 7 - - - 2.9 -

183

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 301 Phylloscartes ventralis 10 115 5.2 53.4 2.6 56.6 3.3 17.7 1.6 12.7 1 302 Phylloscartes virescens 2 114 11.3 54 2.8 56 2.8 16.5 1.1 11.9 1.1 303 Pitangus sulfuratus * 11 222.5 12.4 117.7 6.4 97.5 7.9 25.3 2 24.7 1.8 304 Platyrinchus cancrominus 10 90 5 54.7 3.2 31 3.7 15.8 0.4 12.9 1.2 305 Platyrinchus coronatus 10 84.4 4.7 53.5 3.8 26.3 1.9 13.1 0.6 11.8 1.2 306 Platyrinchus flavigularis 11 99.8 3.1 62.2 1.5 33 2.6 13.1 0.4 12.9 0.7 307 Platyrinchus leucoryphus 2 122.5 16.3 74 0 48 1.4 15.6 0.4 14 0.3 308 Platyrinchus mystaceus 10 87.9 6.5 52 2.4 30.8 4 16.1 0.5 12.6 1.2 309 Platyrinchus platyrhynchos 10 100.4 5.7 62.9 2.3 36.2 1.8 12.9 0.8 12.3 0.8 310 Platyrinchus saturatus 10 90.6 5 55.9 3.2 31.4 3.5 16.5 1.1 13.8 0.9 311 Poecilotriccus albifacies 1 88 - 47 - 40 - 16.6 - 11.2 - 312 Poecilotriccus capitalis 10 90.1 4.1 46.5 1.4 34.8 3.1 16.3 1.3 12 0.9 313 Poecilotriccus ruficeps 9 93.2 4.4 46.8 1.3 40.4 3.7 16.6 0.7 13 0.8 314 Polioxolmis rufipennis 2 198 8.5 125.5 3.5 93.5 0.7 27.4 0.4 20.9 0.1 315 Polystictus pectoralis 10 93.9 6 41.1 5.5 42.3 5.8 16.7 1.1 12.7 0.9 316 Polystictus superciliaris 0 ------317 Pseudelaenia leucospodia 10 110.5 3.8 59.5 5.5 54.4 4.1 19.2 1.2 13.8 0.5 318 Pseudocolopteryx acutipennis 11 102 4.4 44.8 1.8 45 3.6 18.2 0.6 15 1.1 319 Pseudocolopteryx dinellianus 1 118 - 47 - 60 - 19 - 14.1 - 320 Pseudocolopteryx flaviventris 10 113.8 4.2 47.3 2.3 52.3 1.8 18.7 0.7 16 0.8 321 Pseudocolopteryx pelzelni 10 108.8 3.2 56 2.8 47.2 3.9 19.9 0.6 15.3 0.8 322 Pseudocolopteryx sclateri 4 104 3.7 40.3 1.3 46 0.8 17.6 0.6 15 1.4 323 Pseudotriccus pelzelni 0 ------324 Pseudotriccus ruficeps 0 ------325 Pseudotriccus simplex 9 103.2 4.6 54.2 2.8 46.2 3.7 19 0.9 14.9 0.8 326 Pyrocephalus rubinus * 10 130.1 3.8 81.3 2.3 63.1 2.8 15.7 0.9 14.1 0.6 327 Pyrrhomyias cinnamomea 10 125 3.8 68.3 3.4 61.2 0.8 12.5 0.8 11.9 0.8 328 Ramphotrigon fuscicauda 1 154 - 76 - 82 - 18.2 - 13.4 - 329 Ramphotrigon megacephala 2 132 2.8 60 0 61 1.4 15.3 1.1 12.5 0.1 330 Ramphotrigon ruficauda 10 150.1 8.9 76.4 3 72.1 2.8 15.8 0.9 13.5 1.2

184

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 301 Phylloscartes ventralis 10 12.9 0.8 6.7 0.4 3.3 0.2 3.2 0.2 302 Phylloscartes virescens 2 13.1 1.1 7.1 1.1 3.4 0.1 3.3 0.2 303 Pitangus sulfuratus * 11 33.6 2.5 17 0.9 9.9 1 9.7 0.6 304 Platyrinchus cancrominus 10 14.4 0.7 10 0.6 7.4 0.6 3.3 0.3 305 Platyrinchus coronatus 10 11 2.2 7.8 1.4 6.4 0.5 3.3 0.4 306 Platyrinchus flavigularis 11 12.3 1.2 10.1 2 8.3 0.4 4 0.3 307 Platyrinchus leucoryphus 2 15.9 0.4 14.6 1.3 10 - 4.5 0.3 308 Platyrinchus mystaceus 10 13.7 0.5 9.1 0.8 6.7 0.8 3.3 0.3 309 Platyrinchus platyrhynchos 10 14.4 1 12.9 0.7 10 0.5 4.4 0.3 310 Platyrinchus saturatus 10 14.2 0.9 10.8 0.7 8.4 0.4 3.7 0.4 311 Poecilotriccus albifacies 1 13.4 - 8.2 - - - 2.8 - 312 Poecilotriccus capitalis 10 13.6 0.5 8 0.6 5.2 0.3 3.5 0.3 313 Poecilotriccus ruficeps 9 14.1 0.8 6.7 0.5 4.3 0.3 3.2 0.3 314 Polioxolmis rufipennis 2 25.7 1.3 11.3 0.1 6 - 5.3 0.1 315 Polystictus pectoralis 10 11.8 0.6 6.9 0.8 3.6 0.5 2.9 0.2 316 Polystictus superciliaris 0 ------317 Pseudelaenia leucospodia 10 12.8 0.6 8.1 0.6 3.8 0.3 3.4 0.3 318 Pseudocolopteryx acutipennis 11 14 0.6 6.5 0.4 3.3 0.2 3.1 0.3 319 Pseudocolopteryx dinellianus 1 13.3 - 9.2 - 3.5 - 2.9 - 320 Pseudocolopteryx flaviventris 10 14.6 0.9 7 0.6 3.8 0.1 3 0.4 321 Pseudocolopteryx pelzelni 10 13.6 1 8.2 0.4 4.4 0.3 3.7 0.2 322 Pseudocolopteryx sclateri 4 12.3 0.6 6.4 0.4 3.4 0.3 3.1 0.2 323 Pseudotriccus pelzelni 0 ------324 Pseudotriccus ruficeps 0 ------325 Pseudotriccus simplex 9 13.1 0.5 7.7 0.5 3.8 0.3 3.4 0.2 326 Pyrocephalus rubinus * 10 16.7 1.1 8.3 0.4 5.6 0.3 4.3 0.3 327 Pyrrhomyias cinnamomea 10 14.4 0.7 8.6 0.3 6 0.3 3.3 0.2 328 Ramphotrigon fuscicauda 1 18.9 - 10.9 - 6.5 - 4.6 - 329 Ramphotrigon megacephala 2 16 0 10.7 0.5 10.4 6.1 3.9 0.1 330 Ramphotrigon ruficauda 10 17.7 0.9 11.3 0.6 6.7 0.7 4.7 0.2

185

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 331 Rhynchocyclus brevirostris 10 150.1 6.9 76.2 3.7 71.8 4.8 17.9 0.6 14.6 0.9 332 Rhynchocyctus fulvipectus 4 151.5 10.5 74.3 3.9 66.5 5.4 18.2 1.7 15.3 0.7 333 Rhynchocyctus olivaceus 10 143.1 5.7 72.3 2.1 66 3.9 17.2 1.8 15.2 1.4 334 Rhytipterna holerythra * 5 189.4 5.9 103.8 1.9 96.4 4.1 23.8 1.2 18.9 0.7 335 Rhytipterna immunda 0 ------336 Rhytipterna simplex * 6 194.8 8.9 94.3 4.4 95.8 4 21.3 0.9 17.2 1.2 337 Satrapa icterophrys 10 160.6 8.6 88.3 3.2 74.7 3.7 19.1 0.7 17.5 0.8 338 Sayornis nigricans * 10 167 8.2 90.2 4.1 81.8 4.2 17.4 1.1 15.4 0.4 339 Sayornis phoebe * 10 158.3 5.2 85 2.5 74 3.3 17 0.8 14.9 1.2 340 Sayornis saya * 10 178.6 5.6 102.9 5.2 88.4 5.1 20.1 1.3 17.3 0.5 341 Serpophaga cinerea 12 104.4 4.5 53.5 2.8 47.5 3.3 16.6 0.9 12.8 0.6 342 Serpophaga hypoleuca 3 108.7 7.8 48.3 1.2 49 2 16.7 0.5 11.8 1.3 343 Serpophaga munda 10 105.5 4.6 48.7 2.4 51.1 4.5 17 1.2 11.8 0.8 344 Serpophaga nigricans 10 116.7 3.4 55.7 1.3 54.3 3.2 17.4 1.1 13.9 0.9 345 Serpophaga subcristata 12 104.8 7.3 46.8 2.3 48.8 4.1 16.2 0.6 11.3 0.6 346 Silvicultrix diadema 10 115 4.6 60 2.9 50.3 2.3 17.9 0.8 13.8 0.9 347 Silvicultrix frontalis 6 121.2 9.1 65.7 2.8 55.2 4 20.9 1.4 14.5 1 348 Silvicultrix jelskii 1 124 - 62 - 59 - 18.7 - 13.8 - 349 Silvicultrix pulchella 1 112 - 65 - 55 - 20 - 14.5 - 350 Sirystes sibilator 10 184.9 6 96.2 3.2 88.7 3.6 18.9 0.7 17.6 0.9 351 Stigmatura budytoides 10 140.5 2.3 59.9 3.9 77.4 2.8 21.2 0.5 13.3 0.7 352 Stigmatura napensis 9 121.2 3.9 52.1 1.8 62.9 6.1 18.1 1.5 12.8 1.1 353 Sublegatus arenarum 0 ------354 Sublegatus modestus * 11 126.7 6.7 68.2 2.8 64.8 2.4 17.5 0.9 13.2 1 355 Sublegatus obscurior * 1 144 - 73 - 70 - 18.4 - 12.9 - 356 Suiriri affinis 1 144 - 83 - 75 - 20 - 14.2 - 357 Suiriri suiriri 10 141.4 2.7 73 1.4 67.8 1.6 19.1 0.7 13.5 0.5 358 Tachuris rubigastra 10 101.9 4.7 48.6 1.9 42.6 2.7 18.9 0.7 17.5 0.6 359 Teniotriccus andrei 0 ------360 Tityra cayana 1 215 - 123 - 73 - 24 - 21 -

186

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 331 Rhynchocyclus brevirostris 10 17 1.1 13.5 1 8.1 0.9 5.1 0.5 332 Rhynchocyctus fulvipectus 4 17.8 1.1 14.9 1.4 9 0.3 5.5 0.6 333 Rhynchocyctus olivaceus 10 16.8 0.6 13.9 1.2 9 1 5.5 0.3 334 Rhytipterna holerythra * 5 24 1.1 13.7 0.8 7.3 0.6 6 0.4 335 Rhytipterna immunda 0 ------336 Rhytipterna simplex * 6 23.4 1.5 13.4 0.6 7.4 0.6 6.3 0.3 337 Satrapa icterophrys 10 17.9 1.1 9.9 1 4.8 0.3 4 0.4 338 Sayornis nigricans * 10 19 1.1 8.9 0.9 5.6 0.6 4.6 0.4 339 Sayornis phoebe * 10 18.7 1.1 9.1 0.6 5.9 0.4 4.6 0.4 340 Sayornis saya * 10 20.5 1.4 9.6 0.9 6.2 0.4 4.6 0.5 341 Serpophaga cinerea 12 12.3 0.7 6.4 0.6 3.4 0.3 2.9 0.3 342 Serpophaga hypoleuca 3 12.2 0.4 6.7 0.4 3.4 0.1 2.9 0.2 343 Serpophaga munda 10 11.4 0.5 5.5 0.7 3.1 0.3 2.7 0.3 344 Serpophaga nigricans 10 13.8 0.5 6.4 0.8 3.4 0.3 2.9 0.2 345 Serpophaga subcristata 12 11.1 0.6 6 0.7 3 0.3 2.8 0.3 346 Silvicultrix diadema 10 13.1 0.5 7.4 0.5 4.3 0.2 3.3 0.2 347 Silvicultrix frontalis 6 13.1 0.2 7.4 0.3 3.8 0.2 3.1 0.2 348 Silvicultrix jelskii 1 13.5 - 6.3 - 3.6 - 3 - 349 Silvicultrix pulchella 1 13 - 6.5 - 3.7 - 3.3 - 350 Sirystes sibilator 10 22.4 0.7 14.1 0.9 8.6 0.4 6.8 0.6 351 Stigmatura budytoides 10 13 0.7 7.6 0.5 3.3 0.2 3.2 0.2 352 Stigmatura napensis 9 12.9 0.3 7.4 0.3 3.5 0.2 3.2 0.3 353 Sublegatus arenarum 0 ------354 Sublegatus modestus * 11 11.6 0.7 8.2 0.4 4.4 0.2 3.5 0.3 355 Sublegatus obscurior * 1 12.9 - 10.1 - - - 3.7 - 356 Suiriri affinis 1 16.6 - 11.3 - - - 4.1 - 357 Suiriri suiriri 10 14.6 1.1 8.8 0.7 4.7 0.2 4.3 0.3 358 Tachuris rubigastra 10 14.1 0.7 5.7 0.5 2.4 0.3 2.5 0.2 359 Teniotriccus andrei 0 ------360 Tityra cayana 1 27.5 - 19.7 - 10.3 - 10.5 -

187

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 361 Tityra inquisitor 1 167 - 104 - 64 - 20 - 19.5 - 362 Tityra semifasciata 1 204 - 129 - 81 - 26.3 - 25.2 - 363 Todirostrum calopterum 8 86.6 4.3 47.5 1.9 31.6 2 16 0.8 12.5 0.9 364 Todirostrum chrysocrotaphum 10 85.8 3.3 41.1 1.7 30.7 2.1 15.3 0.9 12.3 0.7 365 Todirostrum cinereum 10 93.8 7.4 41.5 1.4 34.5 3.6 18.2 0.4 12 0.8 366 Todirostrum fumifrons 4 88 2.2 42.5 1.7 35.5 1.3 17.2 1.3 11.7 0.7 367 Todirostrum latirostre 10 92.8 4.9 45.9 2.1 37 3 18.2 1.1 12.7 0.8 368 Todirostrum maculatum 10 90.9 3.1 43.9 1.4 35.2 2.6 17 1.3 12.2 0.9 369 Todirostrum nigriceps 10 78.4 4.3 38.5 1.5 27.4 2.1 14.7 0.8 11 1 370 Todirostrum pictum 5 93.6 0.5 41 1.2 33.8 2.3 16.5 1 11.6 0.7 371 Todirostrum plumbeiceps 10 94.3 5.9 42.9 2.1 39.8 3.2 18.3 0.9 13.3 1.1 372 Todirostrum poliocephalum 7 91.3 5.4 41.6 1.4 39.3 2.1 16.8 0.9 11.4 0.5 373 Todirostrum pulchellum 0 ------374 Todirostrum russatum 0 ------375 Todirostrum senex 0 ------376 Todirostrum sylvia 10 93.2 3.7 46.6 1 35.2 1.9 18.1 0.4 13 0.7 377 Todirostrum viridanum 0 ------378 Tolmomyias assimilis 10 123.5 2.6 63.3 3.2 55.7 2.8 16.4 0.5 12.3 0.9 379 Tolmomyias flaviventris 9 117.7 3.2 58.1 2.2 53.7 4 16.9 0.9 12.2 0.4 380 Tolmomyias poliocephalus 10 109.4 6.1 53.5 2.1 48.9 2.2 15.3 0.5 10.9 1 381 Tolmomyias sulphurescens 10 131.2 7.8 63.8 3.2 61.7 4 18.4 0.9 12.3 1.2 382 Tyrannopsis sulphurea 10 194 8 108.9 3.1 86.1 5.3 19.6 0.8 21.3 1.3 383 Tyrannulus elatus* 10 99.8 3.2 50.6 2 43.5 3 12.8 0.8 10.1 0.6 384 Tyrannus albogularis * 10 197.6 5.8 101.5 4 95.1 3.8 16.3 0.8 17.9 0.7 385 Tyrannus caudifasciatus * 10 212.7 13.6 107.6 4.9 94.6 6.9 21.7 1.3 21.6 1.2 386 Tyrannus couchii * 10 216.2 5.9 124.8 7.1 105 6.8 18.9 0.7 19.8 1.1 387 Tyrannus crassirostris * 9 224.2 7.2 129.1 2.7 107 4.3 19.3 0.8 20.5 1.2 388 Tyrannus cubensis * 10 253.7 10 130.9 4.3 103.5 4.9 22.4 1.4 25.2 0.9 389 Tyrannus domenicensis * 10 215.3 4.4 115.5 3.7 99.5 4.5 18 0.7 19.7 0.9 390 Tyrannus fortificata * 10 342.9 18.6 124.4 3.5 238.1 21.7 18.2 0.6 18.6 0.9

188

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 361 Tityra inquisitor 1 27 - 16.5 - 10.4 - 10.4 - 362 Tityra semifasciata 1 29.7 - 22.2 - 11.2 - 10.7 - 363 Todirostrum calopterum 8 13.4 0.5 7.5 0.7 4.9 0.2 3.3 0.2 364 Todirostrum chrysocrotaphum 10 14.6 0.6 8 0.3 5.2 0.9 3.3 0.2 365 Todirostrum cinereum 10 16.5 1.1 6.3 0.4 4.5 0.1 3.1 0.3 366 Todirostrum fumifrons 4 14.8 0.4 7.5 1.1 4.7 0.5 2.9 0.2 367 Todirostrum latirostre 10 15.1 0.6 7.7 0.7 4.6 0.5 3.2 0.3 368 Todirostrum maculatum 10 17 0.8 7.7 0.6 4.8 0.2 3.2 0.2 369 Todirostrum nigriceps 10 14.3 0.3 7.3 0.6 5.4 0.2 3 0.2 370 Todirostrum pictum 5 15.8 0.4 7.2 0.5 5.4 0.4 3.2 0.1 371 Todirostrum plumbeiceps 10 14.2 0.7 6.9 0.7 3.8 0.4 2.9 0.2 372 Todirostrum poliocephalum 7 13.9 0.9 6.6 0.8 4.5 0.3 2.9 0.4 373 Todirostrum pulchellum 0 ------374 Todirostrum russatum 0 ------375 Todirostrum senex 0 ------376 Todirostrum sylvia 10 15.1 0.6 7.9 0.4 5.2 0.3 3.3 0.2 377 Todirostrum viridanum 0 ------378 Tolmomyias assimilis 10 14.4 0.8 10.5 0.6 5.4 0.4 3.9 0.3 379 Tolmomyias flaviventris 9 13.3 0.5 10.3 0.6 5.5 0.5 4 0.3 380 Tolmomyias poliocephalus 10 13.1 0.9 9.6 0.9 5.1 0.6 3.6 0.3 381 Tolmomyias sulphurescens 10 15.1 0.7 11.8 0.6 5.3 0.5 4.1 0.4 382 Tyrannopsis sulphurea 10 21.8 0.5 16 1.3 9.2 0.8 6.9 0.5 383 Tyrannulus elatus* 10 9.4 0.5 5.1 0.6 - - 3.2 0.2 384 Tyrannus albogularis * 10 23.7 0.8 15.2 1.1 9.3 0.8 6.4 0.5 385 Tyrannus caudifasciatus * 10 29.2 1.6 15.9 1.4 9.1 0.6 7.2 0.5 386 Tyrannus couchii * 10 26.1 1.5 16.9 1.1 10.9 0.5 7.9 0.4 387 Tyrannus crassirostris * 9 29.4 1.1 19 2.4 12.9 0.6 10.5 0.2 388 Tyrannus cubensis * 10 38.2 1.2 20.4 1.2 14.6 0.8 11.8 0.4 389 Tyrannus domenicensis * 10 30.1 0.8 15.7 0.7 11.1 0.5 8.6 0.4 390 Tyrannus fortificata * 10 23.3 0.7 13.8 0.9 8.3 0.5 6.4 0.3

189

Appendix I. Continued Species N Total SD Wing SD Tail SD Tarsus SD Toe SD 391 Tyrannus melancholicus * 10 196.8 3.9 112.1 3.6 93.3 5.9 17.7 0.9 19.1 0.9 392 Tyrannus niveigularis * 2 170.5 4.9 96.5 14.8 81.5 7.8 19 1.8 17.1 0.1 393 Tyrannus savana * 7 359.6 44.4 108.6 4.9 262.3 44.7 17 2.2 17.6 0.4 394 Tyrannus tyrannus * 20 197.2 8.6 117.1 4.3 88 5.3 18.5 1.1 18.5 0.9 395 Tyrannus verticalis * 20 205.1 5.8 128.6 5 99.6 4.9 18.3 0.7 19.5 1.2 396 Tyrannus vociferans * 20 208.8 5.1 128.5 4.2 97 3.9 19.2 0.8 19.7 1.1 397 Uromyias agilis * 9 133.7 5.5 57.6 2.6 70.8 1.9 18.6 0.7 14.6 0.9 398 Uromyias agraphia * 10 126.2 4.2 56.8 2.6 67.5 4.2 18.2 1.1 14.3 0.6 399 Xenopsaris albinucha 1 123 - 64 - 59 - 13.7 - 11.2 - 400 Xenotriccus callizonus 1 126 - 58 - 59 - 17.1 - 12.9 - 401 Xenotriccus mexicanus 1 136 - 61 - 66 - 18.1 - 13 - 402 Xolmis cinerea 10 215.7 8.5 137.2 3.9 98.9 7.9 28.6 1.3 25.4 1.6 403 Xolmis coronata 9 212.2 7.8 124.2 7.2 98.4 4.4 29.2 0.7 22.6 1.4 404 Xolmis irupero 10 176 8.8 105.9 5 83.2 4.1 23.3 0.6 19.5 1 405 Xolmis pyrope 10 199.2 9.3 107.8 3.9 94.1 6.1 26.3 1 22.1 1.6 406 Xolmis rubetra 5 181.2 3.4 114 6.3 81.4 4.8 27.9 2.4 20.9 2.1 407 Xolmis salinarum 0 ------408 Xolmis velata 6 197.8 6.6 120.3 3.9 91 5.2 26.5 0.8 22.9 1.4 409 Zimmerius bolivianus 10 115.1 5.8 59.5 3.7 57.1 3.2 16.7 0.9 12.5 0.9 410 Zimmerius chrysops 10 108.2 8.7 54.2 4.1 50.4 4.3 16.1 0.9 11.6 1.1 411 Zimmerius cinereicapillis 2 122 1.4 57 2.8 56.5 3.5 15.6 0.6 12 0.7 412 Zimmerius gracilipes 11 90.7 7.3 44.8 2.9 40.3 4.2 13.9 0.9 10.5 0.9 413 Zimmerius improbus 0 ------414 Zimmerius vilissimus 10 104 6.7 53.2 5.1 50.6 6.9 16.4 0.9 11.8 1.5 415 Zimmerius viridiflavus 9 102.4 5.7 51.9 2.8 46.8 3.3 16.2 0.5 12.2 1.2

190

Appendix I. Continued Species N BL SD BW SD BW2 SD BD SD 391 Tyrannus melancholicus * 10 25.5 1.3 16 0.9 10.1 0.6 7.9 0.4 392 Tyrannus niveigularis * 2 22.5 1.6 13.3 1.8 8.9 0.8 6 0.3 393 Tyrannus savana * 7 18.3 0.6 12.5 0.6 7.1 0.8 6 0.7 394 Tyrannus tyrannus * 20 21.3 1.1 13.5 0.8 8.4 0.5 6.7 0.4 395 Tyrannus verticalis * 20 23.5 1.4 14.8 0.9 8.6 0.4 7 0.5 396 Tyrannus vociferans * 20 24.2 1.2 15.5 1.2 9.2 0.6 7.1 0.4 397 Uromyias agilis * 9 13.4 0.3 7.5 0.6 3.7 0.4 3.6 0.4 398 Uromyias agraphia * 10 13.2 0.8 6.9 0.5 3.7 0.3 3.5 0.2 399 Xenopsaris albinucha 1 12.3 - 9.5 - 4.1 - 3.5 - 400 Xenotriccus callizonus 1 13.9 - 6.9 - - - 3.3 - 401 Xenotriccus mexicanus 1 15.8 - 8.2 - - - 3.9 - 402 Xolmis cinerea 10 25.9 0.8 15.2 1.2 7.2 0.3 6.5 0.5 403 Xolmis coronata 9 23.4 1.6 14.2 1.6 6.6 0.4 5.7 0.3 404 Xolmis irupero 10 21.8 0.7 11.6 0.9 5.7 0.4 4.9 0.4 405 Xolmis pyrope 10 22.6 1.1 13.1 0.6 5.8 0.4 5 0.5 406 Xolmis rubetra 5 21.1 1.1 12.3 0.6 5.3 0.4 4.9 0.3 407 Xolmis salinarum 0 ------408 Xolmis velata 6 24.8 0.8 12.9 0.8 6.7 0.2 5.9 0.3 409 Zimmerius bolivianus 10 11.3 0.5 7 0.5 3.5 0.2 3.4 0.1 410 Zimmerius chrysops 10 10.3 0.6 6.3 0.4 3.4 0.2 3.2 0.3 411 Zimmerius cinereicapillis 2 11.1 0 7.2 0.5 4 - 3.8 0.3 412 Zimmerius gracilipes 11 9.6 0.5 6.3 0.8 3.4 0.3 3.3 0.2 413 Zimmerius improbus 0 ------414 Zimmerius vilissimus 10 10.9 0.6 6.7 0.5 3.4 0.2 3.5 0.2 415 Zimmerius viridiflavus 9 10.5 0.4 6.4 0.5 3.4 0.2 3.3 0.1

191

Appendix II. Species and morphological data used in the CDF, UPGMA and ecological axis analyses.

Lineage Species TOTW TAIL TARTOE BL BW1BW2 BD Elaeniinae 1 Elaenia martinica 153 78 74 20.2 15.1 15.2 11.4 4.2 4.3 2 Elaenia flavogaster 156 78 73 18.6 15.3 14.3 11.3 5.1 4.1 3 Elaenia spectabilis 171 88 85 21.0 15.5 15.6 10.8 4.7 4.3 4 Elaenia albiceps 140 75 68 18.5 14.5 13.7 8.9 4.2 3.9 5 Elaenia parvirostris 134 69 65 17.5 14.0 13.4 8.6 3.4 3.5 6 Elaenia strepera 150 82 71 16.4 14.0 13.3 10.6 4.7 3.8 7 Elaenia mesoleuca 148 75 69 16.7 13.8 13.8 9.3 3.9 3.8 8 Elaenia gigas 172 90 83 18.5 16.6 15.1 11.7 5.5 4.8 9 Elaenia pelzelni 172 89 82 21.3 17.0 16.7 12.2 5.6 4.5 10 Elaenia cristata 134 67 63 17.9 13.2 14.2 9.0 4.8 4.4 11 Elaenia ruficeps 112 57 49 22.3 16.5 13.3 7.6 3.8 3.5 12 Elaenia chiriquensis 140 75 68 16.8 13.4 13.6 9.4 4.4 3.7 13 Elaenia frantzii 149 78 73 17.3 14.4 14.0 8.8 4.5 4.0 14 Elaenia obscura 163 82 83 19.6 15.7 14.9 9.9 4.5 4.6 15 Elaenia pallatangae 140 74 71 16.9 14.1 13.7 7.8 4.1 3.9 16 Elaenia fallax 145 67 71 18.3 13.8 12.8 7.4 3.8 3.5 17 Capsiempis flaveola 110 48 51 16.0 11.1 12.2 7.1 3.6 3.6 18 Mecocerculus leucophrys 142 69 75 22.1 14.6 14.4 8.0 3.7 3.5 19 Mecocerculus poecilocercus 103 56 49 15.9 12.4 11.2 6.1 2.6 3.1 20 Mecocerculus hellmayri 98 57 44 15.8 12.7 11.3 6.3 2.8 3.2 21 Mecocerculus calopterus 107 54 51 16.8 11.7 12.7 6.3 3.1 3.2 22 Mecocerculus minor 117 57 59 16.8 11.7 12.2 6.2 3.0 3.4 192

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 23 Mecocerculus stictopterus 115 61 59 17.4 13.0 12.3 6.6 2.9 3.1 24 Phaeomyias murina 118 61 59 19.8 13.3 12.7 7.8 3.7 3.4 25 Uromyias agilis 134 58 71 18.6 14.6 13.4 7.5 3.7 3.6 26 Uromyias agraphia 126 57 68 18.2 14.3 13.2 6.9 3.7 3.5 27 Anairetes alpinus 131 63 70 20.7 14.8 12.7 6.7 3.1 3.2 28 Anairetes reguloides 125 57 67 20.5 14.1 13.5 6.8 3.0 3.4 29 Anairetes flavirostris 103 49 53 17.9 11.8 11.6 6.4 2.6 2.8 30 Anairetes parulus 105 47 52 18.5 12.1 12.3 6.1 2.6 2.7 31 Camptostoma imberbe 105 54 46 13.9 11.4 11.4 6.6 3.0 3.3 32 Camptostoma obsoletum 107 54 44 14.3 12.0 10.7 6.9 3.3 3.4 33 Euscarthmus meloryphus 100 46 45 19.5 13.6 12.6 6.6 3.1 3.1 34 Euscarthmus rufomarginatus 96 41 50 17.5 13.0 12.9 7.5 3.5 3.3 35 Tyrannulus elatus 105 52 45 13.5 9.5 9.1 6.5 2.4 3.2 36 Myiopagis gaimardii 116 56 57 16.9 12.8 12.7 8.2 3.1 3.5 37 Myiopagis caniceps 123 61 59 16.7 12.7 12.1 7.6 3.6 3.7 38 Myiopagis subplacens 144 69 70 20.0 13.1 14.6 8.2 4.0 4.0 39 Myiopagis flavivertex 122 53 50 15.2 12.3 13.0 8.8 3.4 3.1 40 Myiopagis cotta 135 66 65 16.8 13.1 13.3 7.8 3.7 3.1 41 Myiopagis viridicata 134 64 66 17.1 12.5 13.2 7.9 3.8 3.4 Fluvicolinae 1 Sayornis phoebe 158 85 74 17.0 14.9 18.7 9.1 5.9 4.6 2 Sayornis saya 179 103 88 20.1 17.3 20.5 9.6 6.2 4.6 3 Sayornis nigricans 167 90 82 17.4 15.4 19.0 8.9 5.6 4.6 4 Contopus borealis 177 105 73 14.4 16.0 22.4 13.6 8.7 6.8 5 Contopus pertinax 171 102 86 16.0 14.8 21.5 11.8 8.0 6.1 193

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 6 Contopus lugubris 162 92 80 15.3 14.3 21.1 10.7 7.7 5.2 7 Contopus fumigatus 156 92 71 14.6 13.2 19.7 13.1 7.3 5.1 8 Contopus ochraceus 158 89 76 15.4 14.0 20.9 10.9 8.0 5.1 9 Contopus sordidulus 146 85 69 13.3 12.5 17.7 9.8 6.8 5.0 10 Contopus virens 147 83 68 12.8 12.7 17.7 9.8 6.3 4.8 11 Contopus cinereus 133 71 64 12.7 12.0 16.8 8.8 6.6 4.0 12 Contopus nigrescens 121 65 60 11.1 10.3 14.1 8.3 5.7 3.2 13 Contopus caribaeus 144 73 70 15.2 12.4 20.0 8.7 7.4 4.3 14 Contopus hispanolensis 136 76 70 14.4 12.4 18.0 7.9 6.3 3.7 15 Contopus latirostris 132 66 66 15.0 11.8 17.0 8.5 6.9 3.6 16 Empidonax flaviventris 124 65 54 16.0 12.8 14.5 8.4 5.6 3.8 17 Empidonax virescens 135 72 62 15.3 13.5 16.7 9.1 6.3 4.0 18 Empidonax alnorum 136 68 52 15.8 10.9 15.4 9.5 5.6 3.8 19 Empidonax traillii 131 69 60 16.0 13.9 16.0 8.4 5.8 4.1 20 Empidonax albigularis 120 62 57 16.2 14.1 15.2 7.5 5.9 3.5 21 Empidonax minimus 123 62 59 16.2 12.6 14.5 8.0 5.2 3.8 22 Empidonax hammondii 126 68 60 15.7 12.8 14.3 7.2 4.6 3.7 23 Empidonax wrightii 133 71 64 17.9 13.8 16.5 8.4 5.2 3.9 24 Empidonax oberholseri 136 68 63 17.3 13.6 14.9 8.3 5.0 3.9 25 Empidonax affinis 133 72 66 16.4 12.7 14.8 7.7 4.8 3.7 26 Empidonax difficilis 132 68 63 16.2 13.2 15.5 8.2 5.8 3.8 27 Empidonax occidentalis 123 63 60 17.0 13.0 14.9 8.0 5.3 4.2 28 Empidonax flavescens 125 67 58 16.3 13.3 14.9 8.1 5.5 3.9 29 Empidonax fulvifrons 108 59 51 13.7 11.8 13.0 7.1 4.5 3.4 194

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 30 Empidonax atriceps 116 60 55 15.5 13.2 13.1 6.9 4.7 3.4 31 Ochthoeca cinnamomeiventris 124 67 56 18.1 14.3 14.5 8.3 5.1 3.8 32 Ochthoeca rufipectoralis 133 72 67 18.0 14.3 13.7 7.1 4.2 3.4 33 Ochthoeca fumicolor 152 83 75 22.5 16.7 16.2 9.1 4.7 3.9 34 Ochthoeca oenanthoides 159 85 75 21.7 16.1 17.6 9.9 4.6 3.8 35 Ochthoeca leucophrys 151 77 76 21.6 15.9 17.0 8.8 4.4 3.9 36 Ochthoeca piurae 104 61 43 18.2 13.7 14.8 8.0 3.7 3.4 37 Ochthoeca salvini 134 74 64 17.4 14.7 16.7 9.0 5.1 3.9 38 Agriornis montana 236 135 109 33.1 25.2 30.8 14.5 6.3 6.2 39 Agriornis andicola 261 134 114 38.3 29.7 33.3 17.7 7.5 8.6 40 Agriornis livida 264 130 117 37.0 29.4 35.0 18.2 7.6 8.1 41 Agriornis microptera 240 116 111 32.9 24.8 31.4 17.0 7.3 7.6 42 Agriornis murina 183 99 82 27.5 19.7 22.3 11.5 4.9 4.7 43 Gubernetes yetapa 389 122 267 27.0 25.4 28.2 17.3 10.6 8.6 44 Knipolegus straiticeps 128 57 60 19.0 14.6 14.8 9.2 4.4 3.7 45 Knipolegus hudsoni 148 70 70 20.0 15.1 17.1 10.0 5.2 4.4 46 Knipolegus poecilocercus 142 73 65 17.0 14.7 16.0 9.6 5.2 4.1 47 Knipolegus sigatus 152 85 82 22.8 16.8 17.2 8.9 5.8 4.3 48 Knipolegus cyanirostris 152 79 73 19.7 15.4 16.9 9.7 5.5 4.5 49 Knipolegus poecilurus 123 72 61 17.5 13.8 13.0 7.8 4.6 3.3 50 Knipolegus orenocensis 154 80 75 20.1 16.4 18.9 11.1 5.9 4.8 51 Knipolegus aterrimus 162 84 76 23.0 16.9 19.6 10.3 5.1 4.4 52 Knipolegus nigerrimus 172 102 86 23.8 17.0 18.0 10.5 5.9 5.0 53 Knipolegus lophotes 202 113 91 25.0 19.8 22.8 11.8 5.8 5.5 195

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 54 Machetornis rixosus 187 92 85 29.2 23.1 23.7 11.8 6.1 5.1 55 Fluvicola pica 126 64 53 20.5 17.1 18.0 9.3 5.2 4.0 56 Fluvicola albiventer 130 67 58 20.7 17.3 18.2 8.3 4.9 3.4 57 Fluvicola nengeta 138 74 64 23.0 17.9 19.0 8.1 5.2 3.7 58 Muscigralla brevicauda 107 59 36 25.3 17.4 16.5 8.9 4.3 3.7 59 Pyrocephalus rubinus 130 81 63 15.7 14.1 16.7 8.3 5.6 4.3 60 Sublegatus obscurior 144 73 70 18.4 12.9 12.9 10.1 4.4 3.7 61 Sublegatus modestus 127 68 65 17.5 13.2 11.6 8.2 4.4 3.5 Tyranninae 1 Tyrannus niveigularis 171 97 82 19.0 17.1 22.5 13.3 8.9 6.0 2 Tyrannus albogularis 198 102 95 16.3 17.9 23.7 15.2 9.3 6.4 3 Tyrannus melancholicus 197 112 93 17.7 19.1 25.5 16.0 10.1 7.9 4 Tyrannus couchii 216 125 105 18.9 19.8 26.1 16.9 10.9 7.9 5 Tyrannus vociferans 209 128 97 19.2 19.7 24.2 15.5 9.2 7.1 6 Tyrannus crassirostris 224 129 107 19.3 20.5 29.4 19.0 12.9 10.5 7 Tyrannus verticalis 205 129 100 18.3 19.5 23.5 14.8 8.6 7.0 8 Tyrannus fortificata 343 124 238 18.2 18.6 23.3 13.8 8.3 6.4 9 Tyrannus savana 360 109 262 17.0 17.6 18.3 12.5 7.1 6.0 10 Tyrannus tyrannus 197 117 88 18.5 18.5 21.3 13.5 8.4 6.7 11 Tyrannus domenicensis 215 116 100 18.0 19.7 30.1 15.7 11.1 8.6 12 Tyrannus caudifasciatus 213 108 95 21.7 21.6 29.2 15.9 9.1 7.2 13 Tyrannus cubensis 254 131 104 22.4 25.2 38.2 20.4 14.6 11.8 14 Myiodynastes hemichrysus 199 103 94 18.5 18.5 25.2 16.4 11.1 7.4 15 Myiodynastes chrysocephalus 203 109 100 18.0 18.6 27.2 16.9 11.4 8.4 16 Myiodynastes bairdii 212 114 105 24.2 23.1 30.2 18.1 10.6 8.9 196

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 17 Myiodynastes maculatus 203 104 91 19.5 19.7 28.2 17.1 11.5 8.7 18 Myiodynastes luteiventris 190 113 88 19.1 19.6 25.9 16.6 10.2 8.4 19 Pitangus sulfuratus 223 118 98 25.3 24.7 33.6 17.0 9.9 9.7 20 Attila phoenicurus 189 93 87 19.3 18.6 21.8 12.9 6.5 5.6 21 Attila cinnamomeus 184 91 88 21.8 19.5 23.2 14.5 7.2 6.2 22 Attila torridus 218 95 90 25.6 22.8 27.7 15.8 8.8 6.8 23 Attila citriniventris 175 89 75 21.9 19.8 24.5 15.1 7.5 6.6 24 Attila bolivianus 186 96 91 24.8 22.8 27.0 14.9 8.0 6.3 25 Attila rufus 196 91 83 24.8 21.7 29.3 15.4 7.9 6.6 26 Attila spadiceus 181 88 78 25.3 23.4 27.4 15.6 7.7 6.7 27 Myiozetetes cayanensis 165 86 75 18.5 17.6 17.0 11.0 5.7 5.2 28 Myiozetetes similis 172 92 82 19.3 17.5 17.4 11.5 5.9 5.2 29 Myiozetetes granadensis 166 85 76 17.6 17.6 17.9 11.9 6.4 5.1 30 Myiozetetes luteiventris 139 76 67 14.6 14.0 13.5 9.8 5.3 4.3 31 Megaynchus pitangua 218 122 96 20.5 24.3 36.9 17.9 13.7 10.7 32 Rhytipterna holerythra 189 104 96 23.8 18.9 24.0 13.7 7.3 6.0 33 Rhytipterna simplex 195 94 96 21.3 17.2 23.4 13.4 7.4 6.3 34 Myiarchus apicalis 187 91 92 21.1 17.0 23.0 12.0 8.0 5.8 35 Myiarchus semirufus 173 84 88 20.0 16.9 25.5 13.0 7.0 5.3 36 Myiarchus yucatanensis 170 82 85 21.6 16.4 21.5 12.4 7.5 5.9 37 Myiarchus tuberculifer 159 78 75 19.0 15.5 21.4 10.7 7.1 5.2 38 Myiarchus barbirostris 148 72 74 18.0 13.8 19.9 10.1 7.1 4.5 39 Myiarchus swainsoni 178 90 86 19.5 16.7 22.6 10.9 7.4 5.7 40 Myiarchus venezuelensis 183 87 89 21.1 16.0 22.7 12.6 7.6 5.9 197

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 41 Myiarchus panamensis 187 92 93 22.2 16.6 23.9 13.3 8.2 6.1 42 Myiarchus ferox 181 87 90 21.2 17.0 23.5 12.1 7.9 5.8 43 Myiarchus cephalotes 189 90 93 20.3 15.7 21.9 11.9 7.5 5.6 44 Myiarchus phaeocephalus 188 93 92 22.3 16.7 23.8 12.6 7.6 5.6 45 Myiarchus apicalis 187 91 92 21.1 17.0 23.0 12.0 8.0 5.8 46 Myiarchus cinerascens 193 97 98 22.4 17.3 24.1 11.6 7.3 6.4 47 Myiarchus nuttingi 177 91 90 21.3 16.7 22.4 11.6 6.9 5.9 48 Myiarchus crinitis 189 97 93 20.7 18.2 25.4 13.2 8.5 7.1 49 Myiarchus tyrannulus 186 97 93 21.9 18.2 25.0 13.2 8.5 6.6 50 Myiarchus nugator 208 99 96 24.4 18.8 26.8 15.4 8.9 6.7 51 Myiarchus magnirostris 145 68 65 20.7 14.5 19.1 10.2 6.7 4.4 52 Myiarchus validus 209 100 103 23.4 19.7 26.4 15.4 9.0 7.0 53 Myiarchus sagrae 172 81 82 19.6 16.0 22.8 12.4 7.5 5.4 54 Myiarchus stolidus 174 81 84 20.4 16.8 22.6 10.9 7.4 5.2 55 Myiarchus antillarum 172 86 84 21.9 16.7 23.3 12.3 7.1 5.6 56 Myiarchus oberi 189 98 90 23.5 18.5 26.6 13.1 7.8 5.9 Muscicapini 1 Empidornis semipartitus ------2 Bradornis pallidus 168 84 75 19.3 19.7 17.1 10.9 4.9 4.3 3 Bradornis infuscatus 173 103 82 26.5 21.1 18.5 12.5 5.2 5.0 4 Bradornis mariquensis 155 84 77 20.8 17.6 17.0 10.3 4.6 4.3 5 Bradornis pumilus 155 83 68 20.6 16.3 15.3 9.2 4.6 4.1 6 Bradornis microrhynchus 140 86 67 21.4 17.8 17.8 10.0 4.6 4.3 7 Dioptrornis chocolatinus 171 89 82 24.0 18.3 17.3 10.6 5.1 4.7 8 Dioptrornis fischeri 167 89 80 22.0 19.5 17.3 10.0 5.0 4.3 198

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 9 Dioptrornis brunneus ------10 Melaenornis edolioides 183 101 102 21.3 18.3 18.8 11.7 5.4 5.1 11 Melaenornis pammelaina 185 109 99 22.2 19.2 19.1 10.7 5.2 4.7 12 Melaenornis ardesiacus ------13 Melaenornis annamarulae ------14 Fraseria ocreata 181 95 83 23.9 18.9 22.5 13.5 5.7 5.6 15 Fraseria cinerascens 152 81 67 18.5 17.9 17.4 10.5 5.7 4.9 16 Sigelus silens 177 87 82 23.5 20.4 19.7 10.5 4.8 4.7 17 Rhinomyias addita ------18 Rhinomyias oscillans ------19 Rhinomyias brunneata 157 79 65 15.8 18.0 17.3 11.4 5.9 4.3 20 Rhinomyias olivacea 135 75 56 18.6 16.9 16.8 10.7 5.8 4.4 21 Rhinomyias umbratilis 147 77 70 16.0 16.7 18.9 10.3 6.8 4.3 22 Rhinomyias ruficauda 152 81 68 16.2 16.0 17.2 10.2 5.5 4.2 23 Rhinomyias colonus ------24 Rhinomyias gularis 144 86 60 21.5 18.3 20.0 10.4 5.9 5.0 25 Rhinomyias insignis ------26 Rhinomyias albigularis ------27 Rhinomyias goodfellowi ------28 Muscicapa striata 144 87 63 14.1 15.1 17.9 9.0 5.0 3.8 29 Muscicapa gambagae 125 72 56 15.1 15.6 16.5 9.4 4.2 3.9 30 Muscicapa griseisticta 122 81 52 12.8 14.6 14.6 10.7 5.7 3.5 31 Muscicapa sibirica 126 80 58 12.1 13.6 13.9 9.6 5.0 2.9 32 Muscicapa dauurica ------199

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 33 Muscicapa randi ------34 Muscicapa segregata ------35 Muscicapa ruficauda 126 77 61 16.0 14.9 15.8 10.7 5.2 3.8 36 Muscicapa muttui 137 73 57 14.1 14.2 17.8 8.5 5.9 3.7 37 Muscicapa ferruginea 114 71 49 12.3 13.3 12.9 8.9 6.0 2.6 38 Muscicapa ussheri 126 81 50 13.7 13.5 12.2 10.5 5.1 3.1 39 Muscicapa infuscata 138 87 54 14.2 12.6 11.8 10.6 5.2 3.7 40 Muscicapa boehmi 127 81 58 17.1 15.5 15.0 8.8 4.3 4.8 41 Muscicapa aquatica 128 62 57 14.3 13.9 15.9 8.3 5.0 4.7 42 Muscicapa olivascens ------43 Muscicapa lendu 137 75 61 15.1 15.5 14.2 9.5 4.8 3.9 44 Muscicapa itombwensis ------45 Muscicapa adusta 116 68 52 14.7 14.5 14.8 8.4 4.8 3.7 46 Muscicapa epulata 104 55 48 - 12.3 13.8 7.9 4.7 3.0 47 Muscicapa sethsmithii 94 55 36 9.8 10.8 11.9 8.0 5.1 3.0 48 Muscicapa comitata 133 65 58 14.5 13.6 14.4 10.1 6.8 3.6 49 Muscicapa tessmanni ------50 Muscicapa cassini 124 67 57 12.8 15.4 17.3 8.8 5.5 3.6 51 Muscicapa caerulescens 133 69 68 15.7 15.0 16.1 8.5 5.0 3.8 52 Myioparus griseigularis 131 67 57 17.5 15.0 15.6 8.5 4.5 4.0 53 Myioparus plumbeus 129 73 66 18.0 13.8 15.3 8.4 4.4 3.7 54 Humblotia flavirostris ------55 Ficedula hypoleuca 125 75 50 17.2 15.5 15.0 8.3 4.9 3.5 56 Ficedula albicollis 132 82 62 16.7 15.7 13.5 8.9 4.1 3.3 200

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 57 Ficedula semitorquata 123 80 57 14.5 15.6 13.0 6.8 4.3 3.0 58 Ficedula zanthopygia 126 70 52 16.5 15.3 15.3 8.7 5.3 3.7 59 Ficedula narcissina 115 74 54 15.0 15.7 14.1 9.3 4.6 4.0 60 Ficedula beijingnica ------61 Ficedula mugimaki 124 73 55 15.0 15.7 14.3 8.2 3.9 3.7 62 Ficedula hodgsonii 123 71 57 15.7 15.7 13.2 8.0 4.1 2.6 63 Ficedula strophiata 138 75 62 18.3 15.8 14.0 7.0 3.8 3.0 64 Ficedula parva 118 69 58 16.2 14.7 14.1 8.5 3.8 3.3 65 Ficedula subrubra ------66 Ficedula monileger 120 61 54 19.4 17.4 13.6 8.5 5.0 3.4 67 Ficedula solitaris 98 57 40 16.0 14.0 12.9 7.6 4.3 2.8 68 Ficedula hyperythra 104 59 43 18.0 16.1 13.6 7.6 4.5 3.0 69 Ficedula dumetoria 109 58 45 15.7 14.9 14.3 7.1 5.2 3.7 70 Ficedula rufigula 116 63 53 18.5 16.0 16.0 8.1 4.3 3.8 71 Ficedula buruensis 124 67 47 19.3 18.4 15.8 8.6 5.2 3.5 72 Ficedula basilanica 113 65 44 21.2 18.9 18.0 9.3 5.0 4.3 73 Ficedula henrici 133 68 48 22.0 17.7 15.7 8.4 4.7 3.6 74 Ficedula harterti ------75 Ficedula platenae 119 60 47 21.1 16.2 14.6 8.7 5.1 3.9 76 Ficedula crypta ------77 Ficedula bonthaina ------78 Ficedula westermanni 105 57 47 14.6 13.8 12.7 6.2 4.0 3.2 79 Ficedula superciliaris 107 68 52 15.4 12.9 14.3 7.7 4.2 3.4 80 Ficedula tricolor 121 61 59 18.2 15.1 12.7 6.0 3.5 2.9 201

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 81 Ficedula sapphira 121 62 50 15.7 14.8 13.7 7.3 3.6 3.2 82 Ficedula nigrorufa 118 62 55 19.7 15.6 14.6 7.8 4.9 3.5 83 Ficedula timorensis ------84 Cyanoptila cyanomelana 153 90 68 16.2 16.2 17.8 10.4 5.5 4.1 85 Eumyias thalassina 169 85 79 16.1 15.4 15.0 9.4 5.5 3.2 86 Eumyias sordida 146 77 62 17.7 17.3 18.2 11.7 6.3 4.1 87 Eumyias panayensis 145 77 65 19.1 17.4 15.5 11.3 6.0 3.2 88 Eumyias albicaudata 143 79 70 19.7 16.8 14.7 9.5 5.8 4.4 89 Eumyias indigo 133 72 67 16.0 17.7 14.2 10.0 5.6 3.3 90 Niltava grandis 195 106 101 22.9 22.0 22.2 12.2 6.0 5.0 91 Niltava macgrigoriae 113 67 53 16.1 14.9 13.1 7.2 3.5 3.3 92 Niltava davidi 157 87 68 19.2 19.4 16.2 11.4 5.5 3.9 93 Niltava sundara 170 87 78 23.6 21.6 18.5 12.0 4.5 4.1 94 Niltava sumatrana ------95 Niltava vivida 181 99 92 18.2 18.9 16.6 10.0 4.9 4.8 96 Cyornis sanfordi ------97 Cyornis hoevelli 156 86 72 19.5 19.0 19.1 13.4 6.1 4.5 98 Cyornis hyacinthinus 152 88 68 21.6 18.4 12.8 11.8 5.7 4.5 99 Cyornis concretus 160 90 66 22.6 21.7 22.3 12.0 6.2 5.9 100 Cyornis ruckii ------101 Cyornis herioti 143 78 64 19.6 19.1 18.7 11.0 6.6 4.8 102 Cyornis hainanus 135 72 63 14.8 15.3 15.0 10.6 5.1 3.8 103 Cyornis pallipes 155 75 64 18.6 17.5 17.5 11.2 6.0 4.2 104 Cyornis poliogenys 131 74 59 18.0 17.7 15.9 9.6 5.0 3.5 202

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 105 Cyornis unicolor 160 82 82 18.5 17.2 18.5 10.7 5.2 3.8 106 Cyornis rubeculoides 140 69 64 15.7 15.7 15.8 10.7 5.2 4.2 107 Cyornis banyumas 131 69 66 16.8 15.8 16.4 9.9 4.7 3.7 108 Cyornis lemprieri 151 81 68 17.7 16.8 17.5 11.2 5.8 4.5 109 Cyornis superbus ------110 Cyornis caerulatus 138 74 58 16.9 15.5 12.5 10.7 5.7 5.1 111 Cyornis turcosus 150 74 66 17.5 17.0 16.3 11.4 5.5 4.8 112 Cyornis tickelliae 137 74 64 16.9 15.9 17.6 10.4 6.0 4.2 113 Cyornis rufigaster 142 77 70 17.8 16.3 18.5 10.3 5.0 4.1 114 Cyornis omissus 138 75 62 19.1 18.0 17.0 10.3 5.7 4.1 115 Muscicapella hodgsoni 89 50 32 14.9 13.3 11.7 5.2 2.7 2.3 116 Culicicapa ceylonensis 110 58 58 13.3 11.7 12.5 7.0 4.0 2.6 117 Culicicapa helianthea 112 57 52 14.5 11.9 14.5 7.5 4.6 3.2 118 Horizorhinus dohrni ------Monarchini 1 Erythrocercus mccallii 115 45 48 14.5 11.2 11.5 7.0 3.6 3.0 2 Erythrocercus holochlorus 94 47 45 15.7 10.7 11.3 6.5 3.3 2.4 3 Erythrocercus livingstonei 103 51 51 16.5 9.6 11.0 7.2 4.2 3.2 4 Elminia longicauda 160 72 98 15.8 10.3 13.7 6.9 4.1 3.0 5 Elminia albicauda 152 68 90 14.5 13.7 12.1 7.0 4.4 3.1 6 Elminia nigromitrata 120 61 59 14.8 13.9 16.7 7.8 4.8 3.2 7 Elminia albiventris ------8 Elminia albonotata 138 65 71 16.0 12.1 13.6 7.4 5.0 3.6 9 Terpsiphone nitens 151 62 78 16.8 14.2 16.7 8.3 5.5 4.5 10 Terpsiphone cyanomelas 141 64 75 15.9 16.9 14.4 8.5 5.2 3.8 203

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 11 Hypothymis helenae 129 64 68 15.8 12.5 13.2 8.6 5.2 3.7 12 Hypothymis coelestis 157 75 73 14.3 14.0 17.3 10.0 6.1 4.0 13 Hypothymis azurea 149 68 72 15.8 13.1 16.1 8.0 5.7 3.9 14 Eutrichomyias rowleyi ------15 Terpsiphone rufiventer 189 80 96 16.2 15.2 22.2 11.5 7.0 5.6 16 Terpsiphone bedfordi ------17 Terpsiphone rufocinerea ------18 Terpsiphone batesi ------19 Terpsiphone viridis 191 89 115 15.3 14.1 19.0 9.0 6.0 4.4 20 Terpsiphone atrochalybea ------21 Terpsiphone mutata 170 77 89 14.7 14.0 17.2 12.0 6.8 4.7 22 Terpsiphone corvina 360 96 275 18.9 16.4 19.4 11.8 7.5 4.9 23 Terpsiphone bourbonnensis 155 73 82 18.7 13.9 15.7 8.0 5.5 3.1 24 Terpsiphone paradisi 329 110 128 18.0 15.9 25.2 12.2 7.6 5.3 25 Terpsiphone atrocaudata 216 92 131 15.5 15.3 21.0 11.3 6.1 6.4 26 Terpsiphone cinnamomea 214 100 106 17.2 14.6 25.0 13.2 8.9 6.7 27 Terpsiphone cyanescens 200 89 91 17.8 16.7 25.9 12.7 9.2 6.3 28 Chasiempsis sandwichensis 146 67 69 25.5 14.9 16.8 7.5 4.5 3.5 29 Pomarea dimidiata ------30 Pomarea nigra ------31 Pomarea iphis 165 87 86 26.5 19.2 20.7 8.8 5.0 4.6 32 Pomarea mendozae ------33 Pomarea whitneyi 199 111 117 32.8 23.4 24.4 11.0 6.5 6.7 34 Mayrornis versicolor ------204

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 35 Mayrornis lessoni 140 64 62 17.7 13.8 14.8 7.5 4.0 3.5 36 Mayrornis schistaceus ------37 Neolalage banksiana 157 78 72 22.2 18.8 20.0 9.5 4.0 4.4 38 Clytorhynchus pachycephaloides 188 91 84 20.8 17.9 21.3 10.8 5.6 7.9 39 Clytorhynchus vitiensis 188 89 91 24.5 21.4 24.4 10.1 5.8 5.8 40 Clytorhynchus nigrogularis ------41 Clytorhynchus hamlini 184 98 81 23.8 19.9 25.1 12.7 5.5 7.1 42 Metabolus rugensis 215 106 94 28.5 20.0 27.8 10.7 5.7 7.7 43 Monarcha axillaris 161 76 94 20.2 14.6 15.2 11.0 5.2 4.2 44 Monarcha rubiensis 175 95 86 19.0 15.0 20.2 11.2 7.2 5.2 45 Monarcha cinerascens 170 82 80 22.7 18.2 24.5 10.7 6.8 6.3 46 Monarcha frater 161 86 77 21.6 15.4 20.8 11.0 6.5 6.3 47 Monarcha melanopsis 164 92 81 18.7 15.0 21.0 10.0 6.4 5.8 48 Monarcha erythrostictus ------49 Monarcha castaneiventris 169 81 85 19.2 13.0 19.2 10.7 6.5 5.6 50 Monarcha richardsii ------51 Monarcha pileatus 127 66 66 18.5 14.4 15.7 8.4 4.6 3.8 52 Monarcha castus ------53 Monarcha leucotis 137 72 69 18.0 14.2 16.5 8.0 4.4 4.3 54 Monarcha guttulus 156 75 80 16.8 14.0 17.0 9.2 5.1 5.0 55 Monarcha mundus 176 87 80 19.9 17.3 21.1 9.3 4.4 5.2 56 Monarcha trivirgatus 146 75 75 16.0 15.4 16.2 9.5 5.7 4.0 57 Monarcha sacerdotum ------58 Monarcha everetti 134 67 70 18.1 12.6 15.8 8.9 4.9 3.7 205

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 59 Monarcha loricatus 176 86 82 22.3 17.6 18.4 8.7 4.8 5.0 60 Monarcha boanensis ------61 Monarcha leucurus 162 82 82 18.5 14.7 17.0 10.5 5.4 4.6 62 Monarcha julianae ------63 Monarcha manadensis 166 87 78 16.7 15.0 17.4 12.4 5.1 4.7 64 Monarcha brehmii ------65 Monarcha infelix 158 82 71 15.0 14.5 17.1 9.0 5.4 5.1 66 Monarcha menckei ------67 Monarcha verticalis 163 91 81 17.8 15.5 17.8 9.6 5.6 5.1 68 Monarcha barbatus 175 80 78 19.5 17.2 17.7 11.0 6.3 4.7 69 Monarcha browni ------70 Monarcha viduus ------71 Monarcha godeffroyi 154 86 77 25.6 20.7 21.2 10.0 6.1 4.7 72 Monarcha takatsukasae 148 67 70 23.5 17.0 18.4 8.4 5.2 3.8 73 Monarcha chrysomela 136 77 67 19.9 14.3 18.3 10.2 5.3 5.0 74 Arses telescophthalmus 153 78 80 15.9 13.0 16.8 9.5 5.6 4.0 75 Arses insularis ------76 Arses kaupi 151 79 83 19.3 17.0 14.9 8.4 4.7 3.5 77 Myiagra freycineti 145 67 60 17.9 14.0 16.2 8.4 6.0 3.4 78 Myiagra erythrops 134 72 58 20.7 13.6 16.6 9.0 6.6 4.2 79 Myiagra oceanica 163 80 80 22.0 15.4 20.7 10.4 8.0 5.1 80 Myiagra pluto 170 83 77 18.9 15.7 18.3 10.6 7.5 4.2 81 Myiagra atra ------82 Myiagra galeata 142 67 65 16.1 12.9 16.5 8.4 6.5 3.3 206

Appendix II (continued) TOT W TAIL TAR TOE BL BW1 BW2 BD 83 Myiagra rubecula 151 79 71 16.2 13.6 16.0 8.9 6.3 4.1 84 Myiagra ferrocyanea 138 67 60 17.8 14.4 16.6 9.9 6.1 4.0 85 Myiagra cervinicauda ------86 Myiagra caledonica 146 80 67 17.2 12.7 18.0 10.0 7.4 4.1 87 Myiagra vanikorensis 151 73 63 18.0 13.8 18.4 8.2 7.0 3.6 88 Myiagra albiventris ------89 Myiagra azureocapilla 147 84 84 21.3 14.9 17.3 8.3 6.6 3.6 90 Myiagra ruficollis 158 78 71 18.0 13.1 17.1 8.4 7.0 3.9 91 Myiagra cyanoleuca 159 88 80 15.8 13.6 18.3 9.7 7.5 4.2 92 Myiagra inquieta 193 108 106 19.5 17.7 24.6 12.7 6.1 4.7 93 Myiagra alecto 176 88 87 19.5 16.8 21.8 10.8 6.6 4.9 94 Myiagra hebetior 162 81 81 19.3 18.0 21.8 10.7 6.6 4.5 95 Lamprolia victoriae 126 75 51 23.7 17.0 17.5 9.0 4.4 3.8 96 Machaerirhynchus flaviventer ------97 Machaerirhynchus nigripectus 132 60 57 17.2 13.0 16.7 9.2 7.5 3.9 98 Grallina cyanoleuca 279 180 133 41.4 28.4 27.2 12.3 5.9 5.8 99 Grallina bruijni 185 106 86 25.7 21.4 23.9 12.3 5.0 5.6

207

Appendix III. Species used in the analyses, lineage membership designations (1 = elaenia, 2 = tody-tyrants), number of specimens measured (n) and mean morphological measurements by species for each variable. *Cnipodectes subrunneus was not used in any subclade designation but was included in the total lineage disparity.

Bill Genus species LineageSubclade n Wing Tail Tarsus Length Width Depth

Elaenia martinica 1 e1 1177.91 73.9120.17 15.24 11.41 4.32 Elaenia spectabilis 1 e1 1188.08 85.1721.04 15.57 10.75 4.28 Elaenia parvirostris 1 e1 1169.36 65.2717.51 13.41 8.62 3.51 Elaenia mesoleuca 1 e1 1175.09 69.1816.65 13.83 9.30 3.82 Elaenia flavogaster 1 e1 1177.64 72.6418.56 14.34 11.30 4.10 Elaenia chiriquensis 1 e1 1375.14 67.9216.82 13.58 9.38 3.70 Elaenia strepera 1 e1 2 82.00 70.5016.35 13.25 10.60 3.80 Elaenia gigas 1 e1 5 89.39 82.7918.27 15.10 11.54 4.75 Elaenia pelzelni 1 e1 1189.45 82.1921.29 16.71 12.17 4.48 Elaenia cristata 1 e1 1367.17 63.2317.89 14.23 9.00 4.42 Elaenia frantzii 1 e1 1478.43 73.0617.33 14.02 8.82 3.99 Elaenia obscura 1 e1 1281.92 82.6619.55 14.86 9.92 4.65 Elaenia pallatangae 1 e1 1073.88 70.8916.92 13.69 7.84 3.87

208

Appendix III Continued Elaenia fallax 1 e1 1066.60 70.8918.33 12.78 7.44 3.52 Elaenia albiceps 1 e1 1475.35 68.2818.46 13.70 8.94 3.85 Myiopagis elatus 1 e1 1050.61 43.5012.75 9.35 5.05 3.15 Myiopagis gaimardii 1 e1 1155.73 56.5516.91 12.66 8.20 3.51 Myiopagis viridicata 1 e1 1264.08 65.4917.05 13.20 7.87 3.40 Myiopagis cotta 1 e1 3 66.67 66.3317.00 13.27 7.77 3.33 Myiopagis caniceps 1 e1 6 60.34 57.8416.52 12.05 7.52 3.70 Myiopagis subplacens 1 e1 1167.19 68.0019.52 14.41 8.23 3.91 Tyranniscus plumbeiceps 1 e1 4 57.50 54.2514.83 10.80 6.07 3.07 Tyranniscus nigrocapillus 1 e1 1062.30 52.9116.05 10.56 6.50 3.20 Tyranniscus cinereiceps 1 e1 1061.00 45.2013.84 11.01 7.02 3.28 Tyranniscus uropygialis 1 e1 1060.60 52.4016.86 10.72 6.57 3.12 Mecocerculus minor 1 e1 1157.45 58.9116.85 12.21 6.25 3.36 Mecocerculus calopterus 1 e1 8 53.75 50.6316.80 12.69 6.33 3.21 Ornithion inerme 1 e2/e4 8 44.50 34.0013.01 10.56 6.38 3.11 Ornithion semiflavum 1 e2/e4 8 46.25 28.1313.90 9.79 6.94 3.49 Ornithion brunneicapillum 1 e2/e4 1 44.00 29.0013.60 10.70 7.40 3.40 Camptostoma imberbe 1 e2/e4 1053.90 46.2013.92 11.38 6.62 3.29 Camptostoma obsoletum 1 e2/e4 1254.00 43.5814.34 10.66 6.86 3.36 Mecocerculus poecilocercus 1 e2/e4 1455.86 49.2215.89 11.19 6.14 3.11 Mecocerculus stictopterus 1 e2/e4 1061.09 58.9017.42 12.32 6.58 3.11 Mecocerculus hellmayri 1 e2/e4 8 56.49 44.3715.76 11.30 6.25 3.16 Suiriri suiriri 1 e2/e4 1173.91 68.4519.15 14.76 9.06 4.30

209

Appendix III Continued Mecocerculus leucophrys 1 e3/e4 1168.55 75.3422.11 14.41 8.00 3.53 Capsiempis flaveola 1 e3/e4 1 48.00 51.2016.00 12.20 7.10 3.60 Phaeomyias murina 1 e3/e4 2 60.60 59.3119.81 12.72 7.89 3.49 Nesotriccus ridgwayi 1 e3/e4 9 54.50 55.0018.75 17.80 9.00 3.55 Serpophaga nigricans 1 e3/e4 1155.37 53.1317.40 13.83 6.10 2.91 Serpophaga hypoleuca 1 e3/e4 9 48.00 49.0016.50 12.00 6.90 2.80 Serpophaga munda 1 e3/e4 6 49.00 51.5016.76 11.48 5.29 2.73 Serpophaga subcristata 1 e3/e4 9 46.84 48.8316.17 11.12 6.01 2.78 Serpophaga cinerea 1 e3/e4 1053.51 47.5016.60 12.25 6.35 2.90 Uromyias agraphia 1 e3/e4 1056.79 67.5018.20 13.21 6.87 3.54 Anairetes agilis 1 e3/e4 2 56.60 70.0018.68 13.40 7.76 3.72 Anairetes alpinus 1 e3/e4 8 62.75 69.7420.67 12.67 6.70 3.22 Anairetes reguloides 1 e3/e4 2 56.79 66.6020.45 13.54 6.80 3.37 Anairetes flavirostris 1 e3/e4 8 48.80 52.6017.92 11.57 6.41 2.79 Anairetes parulus 1 e3/e4 1247.00 51.7218.52 12.28 6.08 2.72 Pseudocolopteryx acutipennis 1 e4 1244.82 45.0018.23 14.04 6.48 3.12 Pseudocolopteryx flaviventris 1 e4 1045.00 51.0019.50 15.60 7.60 2.40 Pseudocolopteryx sclateri 1 e4 5 39.50 46.0017.15 12.23 6.14 2.95 Polystictus pectoralis 1 e4 1142.84 45.0016.47 11.72 6.93 2.86 Rhynchocyclus brevirostris 2 t1 9 76.10 71.9917.83 16.85 13.19 5.12 Rhynchocyclus olivaceus 2 t1 4 71.75 67.0016.35 16.45 13.75 5.32 Rhynchocyclus fulvipectus 2 t1 2 73.50 65.4918.95 17.50 16.05 5.75 Tolomomyias sulphurescens 2 t1 1063.80 61.7018.38 15.08 11.76 4.11

210

Appendix III Continued Tolomomyias poliocephalus 2 t1 4 53.51 48.2514.83 13.27 9.85 3.62 Tolomomyias flaviventris 2 t1 2 59.50 53.5116.80 13.55 10.10 3.72 Tolomomyias assimilis 2 t1 4 61.25 58.0016.27 14.95 10.92 4.07 Platyrhynchus cancrominus 2 t1 5 53.00 30.8015.76 14.00 9.62 3.18 Platyrhynchus mystaceus 2 t1 7 52.57 31.2816.16 13.61 9.13 3.38 Platyrhynchus coronatus 2 t1 5 54.20 26.4012.94 13.04 8.96 3.68 Platyrhynchus flavigularis 2 t1 1 62.00 33.0013.60 13.20 12.60 3.30 Platyrhynchus platyrhynchos 2 t1 1 61.00 35.0011.50 13.80 11.90 4.50 Platyrhynchus leucoryphus 2 t1 1 74.00 49.0015.30 15.60 15.50 4.70 Onychorrhynchus coronatus 2 t1 3 83.68 68.9916.23 25.57 10.73 4.73 Cnipodectes subbrunneus 2 T* 2 80.50 78.0017.25 17.00 13.40 5.45 Poecilotriccus ruficeps 2 t2 9 46.77 40.4516.63 14.11 6.72 3.19 Todirostrum fumifrons 2 t2 3 42.00 35.3316.76 14.80 7.47 2.93 Todirostrum plumbiceps 2 t2 1042.90 39.8018.29 14.19 6.87 2.92 Todirostrum latirostrecaniceps 2 t2 1045.24 37.8818.60 15.01 7.65 3.18 Todirostrum sylvia 2 t2 1046.60 35.2018.13 15.13 7.87 3.26 Todirostrum albifanes 2 t2 1 47.00 40.0016.60 13.40 8.20 2.80 Todirostrum capitale 2 t2 1046.51 34.8016.27 13.60 8.03 3.48 Todirostrum calopterum 2 t2 8 47.50 31.6216.00 13.40 7.46 3.30 Todirostrum chrysocrotaphum 2 t2 1041.10 30.7015.30 14.56 8.00 3.28 Todirostrum nigriceps 2 t2 4 38.25 30.7515.15 14.28 7.55 2.98 Todirostrum cinereum 2 t2 1041.50 34.5018.22 16.53 6.33 3.12 Todirostrum maculatum 2 t2 1043.90 35.2017.03 16.97 7.67 3.20

211

Appendix III Continued Todirostrum poliocephalum 2 t2 4 41.00 38.2516.35 14.50 7.02 3.10 Todirostrum pictum 2 t2 1 42.00 31.0016.40 16.00 6.70 3.20 Myiornis auricularis 2 t2 2 37.50 33.0013.85 11.90 6.50 3.00 Myiornis albiventris 2 t2 9 37.11 26.8913.29 11.52 6.26 2.73 Myiornis ecaudatus 2 t2 1231.83 16.1711.54 11.48 6.16 2.78 Myiornis atricapillus 2 t2 1035.40 16.3012.67 11.67 6.74 2.79 Hemitriccus minimus 2 t2 6 46.51 36.8314.30 13.95 7.57 3.00 Hemitriccus nidipendulus 2 t2 2 45.00 43.0018.40 13.95 7.55 3.20 Hemitriccus obsoletus 2 t2 2 56.49 54.0020.25 14.10 8.15 3.70 Hemitriccus flammulatus 2 t2 3153.26 47.0717.60 14.46 8.46 3.44 Hemitriccus straiticollis 2 t2 1849.39 43.5517.43 14.67 8.11 3.61 Hemitriccus johannis 2 t2 5 48.60 41.6016.36 14.04 8.56 3.62 Hemitriccus margaritaceiventer 2 t2 3348.45 45.4918.80 14.50 7.72 3.36 Hemitriccus z. zosterops 2 t2 4 48.00 45.2515.02 13.35 8.33 3.33 Hemitriccus z. viridiflavus 2 t2 1152.46 50.7216.07 13.86 8.36 3.26 Hemitriccus z. grisipectus 2 t2 2649.58 47.9214.88 13.32 8.45 3.15 Hemitriccus minor 2 t2 2650.50 44.5015.17 14.46 6.93 3.10 Hemitriccus orbatus 2 t2 1 50.00 50.0018.70 15.10 7.90 3.40 Hemitriccus diops 2 t2 7 51.29 51.7118.01 13.78 8.03 3.41 Hemitriccus spodiops 2 t2 5 46.40 38.0015.12 13.14 6.62 3.38 Hemitriccus rufigularis 2 t2 1451.57 52.1416.56 14.72 7.97 3.16 Hemitriccus granadensis 2 t2 2346.17 43.0815.95 13.44 6.73 3.08 Hemitriccus cinnamomeipectus 2 t2 6 49.17 48.1618.15 15.00 7.92 3.22

212

Appendix III Continued Lophotriccus pileatus 2 t2 1048.80 38.3015.06 12.68 7.22 3.14 Lophotriccus vitiosus 2 t2 1046.60 39.1014.01 12.26 7.61 3.29 Lophotriccus eulophotes 2 t2 1047.00 42.9014.54 12.27 7.60 3.17 Lophotriccus galeatus 2 t2 6 44.50 40.5015.12 12.13 7.24 3.22 Atalotriccus pilaris 2 t2 4 39.00 36.2516.05 12.15 6.58 3.13 Oncostoma cinereigulare 2 t2 1047.90 41.2014.62 13.08 6.99 3.26 Oncostoma olivaceum 2 t2 9 45.11 37.5614.59 13.53 7.73 3.63

213

Appendix IV. Results of the multivariate ellipse analysis on the CPC scores of Elaenia and Tody-tyrant lineages, and all included subclades. Bolded values are the basic parameters of the ellipse analysis. Subclades e2 and e3 comprise e4. These along with e1 comprise Elaenia. Subclades t1 and t2 comprise Todys.

Group X Y n Var X Var Y COV (XY) Eig1 Sd Maj Se Eig1 Eig2 Sd Min Se Eig2 Elaenia CPC1 CPC2 58 2.7E-02 4.1E-03 -1.3E-04 2.7E-02 1.7E-01 5.1E-03 4.1E-03 6.4E-02 7.6E-04 CPC2 CPC3 58 4.1E-03 2.9E-03 1.7E-03 5.3E-03 7.3E-02 9.8E-04 1.7E-03 4.1E-02 3.2E-04 e1 CPC1 CPC2 27 2.4E-02 1.5E-03 2.3E-03 2.4E-02 1.6E-01 6.6E-03 1.3E-03 3.6E-02 3.5E-04 CPC2 CPC3 27 1.5E-02 2.3E-03 4.9E-04 1.5E-02 1.2E-01 4.1E-03 2.3E-03 4.8E-02 6.3E-04 e4 CPC1 CPC2 31 1.5E-02 6.0E-03 -4.5E-03 1.6E-02 1.3E-01 4.2E-03 4.1E-03 6.4E-02 1.0E-03 CPC2 CPC3 31 6.0E-03 2.8E-03 -2.8E-04 6.0E-03 7.8E-02 1.5E-03 2.7E-03 5.2E-02 6.9E-04 e2 CPC1 CPC2 9 2.3E-02 7.3E-03 -9.3E-03 2.7E-02 1.7E-01 1.3E-02 3.0E-03 5.5E-02 1.4E-03 CPC2 CPC3 9 7.3E-03 1.0E-03 1.2E-03 7.5E-03 8.7E-02 3.5E-03 8.1E-04 2.8E-02 3.8E-04 e3 CPC1 CPC2 15 1.2E-02 1.9E-03 -3.7E-04 1.2E-02 1.1E-01 4.3E-03 1.9E-03 4.3E-02 6.9E-04 CPC2 CPC3 15 1.9E-03 1.5E-03 7.1E-04 2.4E-03 4.9E-02 8.9E-04 9.6E-04 3.1E-02 3.5E-04 Todys CPC1 CPC2 57 3.5E-02 7.0E-03 2.0E-04 3.5E-02 1.9E-01 6.6E-03 7.0E-03 8.4E-02 1.3E-03 CPC2 CPC3 57 7.0E-03 1.8E-03 6.0E-05 7.0E-03 8.4E-02 1.3E-03 1.8E-03 4.3E-02 3.4E-04 t1 CPC1 CPC2 14 3.5E-02 7.7E-03 -7.2E-03 3.7E-02 1.9E-01 1.4E-02 5.9E-03 7.7E-02 2.2E-03 CPC2 CPC3 14 7.7E-03 1.3E-03 -2.1E-04 7.7E-03 8.8E-02 2.9E-03 1.3E-03 3.6E-02 4.9E-04 t2 CPC1 CPC2 42 1.4E-02 3.6E-03 -5.6E-03 1.7E-02 1.3E-01 3.6E-03 1.2E-03 3.4E-02 2.6E-04 CPC2 CPC3 42 3.6E-03 2.0E-03 3.6E-04 3.7E-03 6.1E-02 8.0E-04 2.0E-03 4.4E-02 4.3E-04

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Appendix IV continued.

Group X Y m11 m12 m21 m22 Var m11 Se m11 Var m12 Se m12 Shape Se Shape Elaenia CPC1 CPC2 1.0E+00 -5.1E-03 5.1E-03 -1.0E+00 9.4E-08 3.1E-04 3.6E-03 6.0E-02 2.6 3.4E-01 CPC2 CPC3 8.8E-01 4.8E-01 4.8E-01 -8.8E-01 2.7E-03 5.2E-02 9.2E-03 9.6E-02 1.8 2.3E-01 e1 CPC1 CPC2 1.0E+00 9.7E-02 9.7E-02 -1.0E+00 2.0E-05 4.5E-03 2.1E-03 4.6E-02 4.4 8.4E-01 CPC2 CPC3 1.0E+00 3.5E-02 3.5E-02 -1.0E+00 1.0E-05 3.2E-03 7.9E-03 8.9E-02 2.6 4.9E-01 e4 CPC1 CPC2 9.4E-01 -3.4E-01 3.4E-01 -9.4E-01 1.7E-03 4.1E-02 1.2E-02 1.1E-01 2.0 3.6E-01 CPC2 CPC3 1.0E+00 6.3E-02 6.3E-02 -1.0E+00 2.0E-04 1.4E-02 4.9E-02 2.2E-01 1.5 2.7E-01 e2 CPC1 CPC2 9.2E-01 -3.9E-01 3.9E-01 -9.2E-01 2.4E-03 4.9E-02 1.3E-02 1.1E-01 3.0 1.0E+00 CPC2 CPC3 9.9E-01 1.7E-01 1.7E-01 -9.9E-01 4.4E-04 2.1E-02 1.5E-02 1.2E-01 3.1 1.0E+00 e3 CPC1 CPC2 1.0E+00 -3.5E-02 3.5E-02 -1.0E+00 1.8E-05 4.3E-03 1.5E-02 1.2E-01 2.5 6.4E-01 CPC2 CPC3 8.8E-01 4.7E-01 4.7E-01 -8.8E-01 1.6E-02 1.3E-01 5.6E-02 2.4E-01 1.6 4.1E-01 Todys CPC1 CPC2 1.0E+00 6.4E-03 6.4E-03 -1.0E+00 2.2E-07 4.7E-04 5.5E-03 7.4E-02 2.2 3.0E-01 CPC2 CPC3 1.0E+00 1.0E-02 1.0E-02 -1.0E+00 8.3E-07 9.1E-04 8.3E-03 9.1E-02 2.0 2.6E-01 t1 CPC1 CPC2 9.8E-01 -2.2E-01 2.2E-01 -9.8E-01 7.8E-04 2.8E-02 1.5E-02 1.2E-01 2.5 6.7E-01 CPC2 CPC3 1.0E+00 -3.0E-02 3.0E-02 -1.0E+00 1.5E-05 3.9E-03 1.7E-02 1.3E-01 2.4 6.5E-01 t2 CPC1 CPC2 9.2E-01 -3.8E-01 3.8E-01 -9.2E-01 2.9E-04 1.7E-02 1.7E-03 4.1E-02 3.7 5.8E-01 CPC2 CPC3 9.9E-01 1.5E-01 1.5E-01 -9.9E-01 1.3E-03 3.5E-02 5.7E-02 2.4E-01 1.4 2.1E-01

Variables in Table 2: Group = subclade label (see Fig. 1), X = variable 1, Y = variable 2, n = number of species in subclade, Var = variance, COV = covariance, Eig = eigenvalue of singular value decomposition of X and Y, Sd = Standard deviation, Se = standard error.

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