TESTING FOR UNEQUAL AMOUNTS OF EVOLUTION IN A CONTINUOUS CHARACTER ON DIFFERENT BRANCHES OF A PHYLOGENETIC TREE USING LINEAR AND SQUARED‐CHANGE PARSIMONY: AN EXAMPLE USING LESSER ANTILLEAN ANOLIS LIZARDS

Although a large body of work investigating tests of correlated evolution of two continuous characters exists, hypotheses such as character displacement are really tests of whether substantial evolutionary change has occurred on a particular branch or branches of the phylogenetic tree. In this study, we present a methodology for testing such a hypothesis using ancestral character state reconstruction and simulation. Furthermore, we suggest how to investigate the robustness of the hypothesis test by varying the reconstruction methods or simulation parameters. As a case study, we tested a hypothesis of character displacement in body size of Caribbean Anolis lizards. We compared squared‐change, weighted squared‐change, and linear parsimony reconstruction methods, gradual Brownian motion and speciational models of evolution, and several resolution methods for linear parsimony. We used ancestor reconstruction methods to infer the amount of body size evolution, and tested whether evolutionary change in body size was greater on branches of the phylogenetic tree in which a transition from occupying a single‐species island to a two‐species island occurred. Simulations were used to generate null distributions of reconstructed body size change. The hypothesis of character displacement was tested using Wilcoxon Rank‐Sums. When tested against simulated null distributions, all of the reconstruction methods resulted in more significant P‐values than when standard statistical tables were used. These results confirm that P‐values for tests using ancestor reconstruction methods should be assessed via simulation rather than from standard statistical tables. Linear parsimony can produce an infinite number of most parsimonious reconstructions in continuous characters. We present an example of assessing the robustness of our statistical test by exploring the sample space of possible resolutions. We compare ACCTRAN and DELTRAN resolutions of ambiguous character reconstructions in linear parsimony to the most and least conservative resolutions for our particular hypothesis.

[1]  T. Schoener Size Patterns in West Indian Anolis Lizards. II. Correlations with the Sizes of Particular Sympatric Species-Displacement and Convergence , 1970, The American Naturalist.

[2]  Jonathan B. Losos,et al.  Ecomorphology, Performance Capability, and Scaling of West Indian Anolis Lizards: An Evolutionary Analysis , 1990 .

[3]  W. Maddison Squared-Change Parsimony Reconstructions of Ancestral States for Continuous-Valued Characters on a Phylogenetic Tree , 1991 .

[4]  J. Lazell,et al.  The Anoles (Sauria, Iguanidae) of the Lesser Antilles. , 1972 .

[5]  T. Garland,et al.  Why Not to Do Two-Species Comparative Studies: Limitations on Inferring Adaptation , 1994, Physiological Zoology.

[6]  Birgitta Siixén-Tullberg THE EFFECT OF BIASED INCLUSION OF TAXA ON THE CORRELATION BETWEEN DISCRETE CHARACTERS IN PHYLOGENETIC TREES , 1993, Evolution; international journal of organic evolution.

[7]  Theodore Garland,et al.  Phylogenetic Analysis of Covariance by Computer Simulation , 1993 .

[8]  A. Dunham,et al.  THE PARADOX OF THE PHYLOGENY: CHARACTER DISPLACEMENT OF ANALYSES OF BODY SIZE IN ISLAND ANOLIS , 1996, Evolution; international journal of organic evolution.

[9]  E. Williams The Origin of Faunas. Evolution of Lizard Congeners in a Complex Island Fauna: A Trial Analysis , 1972 .

[10]  J. Losos,et al.  A PHYLOGENETIC ANALYSIS OF CHARACTER DISPLACEMENT IN CARIBBEAN ANOLIS LIZARDS , 1990, Evolution; international journal of organic evolution.

[11]  B. Sillén-Tullberg,et al.  EVOLUTION OF GREGARIOUSNESS IN APOSEMATIC BUTTERFLY LARVAE: A PHYLOGENETIC ANALYSIS , 1988, Evolution; international journal of organic evolution.

[12]  E. Martins The Comparative Method in Evolutionary Biology, Paul H. Harvey, Mark D. Pagel. Oxford University Press, Oxford (1991), vii, + 239 Price $24.95 paperback , 1992 .

[13]  J. Felsenstein Phylogenies and the Comparative Method , 1985, The American Naturalist.

[14]  David L. Swofford,et al.  Reconstructing ancestral character states under Wagner parsimony , 1987 .

[15]  R. Huey,et al.  PHYLOGENETIC STUDIES OF COADAPTATION: PREFERRED TEMPERATURES VERSUS OPTIMAL PERFORMANCE TEMPERATURES OF LIZARDS , 1987, Evolution; international journal of organic evolution.

[16]  M. McPeek Testing Hypotheses About Evolutionary Change on Single Branches of a Phylogeny Using Evolutionary Contrasts , 1995, The American Naturalist.

[17]  S. Hedges,et al.  Molecular insights into the relationships and biogeography of West Indian anoline lizards , 1993 .

[18]  Ramón Díaz-Uriarte,et al.  TESTING HYPOTHESES OF CORRELATED EVOLUTION USING PHYLOGENETICALLY INDEPENDENT CONTRASTS: SENSITIVITY TO DEVIATIONS FROM BROWNIAN MOTION , 1996 .

[19]  M. Soulé,et al.  The Relationships of the Puerto Rican Anolis: Electrophoretic and Karyotypic Studies , 1983 .

[20]  T. Garland,et al.  CHAPTER 13 – RECONSTRUCTING ANCESTRAL TRAIT VALUES USING SQUARED-CHANGE PARSIMONY: PLASMA OSMOLARITY AT THE ORIGIN OF AMNIOTES , 1997 .

[21]  P. Martins CONTINUOUS CHARACTERS: A SIMULATION STUDY , 1991 .

[22]  H. Dessauer,et al.  Comparative immunological study of albumins of Anolis lizards of the Caribbean islands , 1981 .

[23]  W. Maddison A METHOD FOR TESTING THE CORRELATED EVOLUTION OF TWO BINARY CHARACTERS: ARE GAINS OR LOSSES CONCENTRATED ON CERTAIN BRANCHES OF A PHYLOGENETIC TREE? , 1990, Evolution; international journal of organic evolution.

[24]  T. Garland,et al.  PHYLOGENETIC ANALYSES OF THE CORRELATED EVOLUTION OF CONTINUOUS CHARACTERS: A SIMULATION STUDY , 1991, Evolution; international journal of organic evolution.

[25]  M. Soulé,et al.  The Relationships of the Anolis crislatellus Species Group: Electrophoretic Analysis , 1980 .