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Toward an Integrative Historical Biogeography1 INTEGR.COMP.BIOL., 43:261±270 (2003) Toward an Integrative Historical Biogeography1 MICHAEL J. DONOGHUE2 AND BRIAN R. MOORE Department of Ecology and Evolutionary Biology, Yale University, P.O. Box 208106, New Haven, Connecticut 06511 SYNOPSIS. Cladistic biogeographic methods remain susceptible to the confounding effects of ``pseudo-con- gruence'' and ``pseudo-incongruence'' because they were not designed to incorporate information on the absolute timing of the diversi®cation of lineages. Consequently, results from cladistic biogeographic studies are dif®cult to interpret and cannot be con®dently attributed to any particular cause. We illustrate these points with concrete examples, paying special attention to recent work on the biogeography of the Northern Hemisphere, and outline ways in which topological and temporal information might be better integrated. The development of historical biogeography over the last few decades provides general insights into the nature of integration through the life of a discipline. INTRODUCTION ing areas of endemism. Willi Hennig (1966) advanced Despite its very promising beginnings, we agree the idea of overlaying phylogenetic trees on geograph- with the assessment of Nelson and Ladiges (1996, p. ic maps to help determine former connections among 54) that cladistic biogeography has yielded few gen- areas and the direction of movement within lineages. uinely new insights over the last twenty years. When Lars Brundin (e.g., 1966) synthesized these two great a research program is unproductive for a long period ideas in his studies of midges in the Southern Hemi- it suggests that something may be wrong with the sphere, searching for general phylogenetic patterns in questions being asked or the methods being used. Here the geographic distribution of multiple clades. Collec- we highlight several elements that have been missing tively, this work represented a signi®cant departure from cladistic biogeographic approaches, focusing par- from (and improvement upon) the previous practice of ticularly on temporal information, and suggest how developing narrative biogeographic scenarios on a these might be integrated. case-by-case basis. The value of incorporating temporal information in In the hands of Donn Rosen, Gareth Nelson, and such studies has already been recognized (see refer- Norman Platnick these insights were subsequently re- ences below). Our aim here is to help characterize and ®ned and formalized (e.g., Rosen, 1975, 1978; Platnick illustrate the problem using concrete examples from and Nelson, 1978; Nelson and Platnick, 1981; Nelson both the Southern and Northern Hemispheres. The and Rosen, 1981). The goal of the resulting cladistic Northern Hemisphere problem provides an especially biogeographic program was to identify general pat- good example of the dif®culties faced by cladistic bio- terns in the relationships among geographical areas of geographic methods, as well as insights into what can endemism, which would therefore suggest the need for be gained by integrating time and directionality. Al- general explanations such as vicariance caused by par- though we do not propose a new biogeographic meth- ticular events in Earth history or concerted dispersal. od, we highlight several possibilities by comparison to In general, this approach entails the compilation of methods recently developed to analyze the coevolution ``taxon area cladograms'' (phylogenies in which the of hosts and their parasites. names of terminal taxa have been replaced by the areas Finally, in the spirit of the symposium that moti- in which they occur) and the search for patterns of vated this paper, we offer some general re¯ections on topological congruence expressed in the form of ``gen- ``integration.'' Speci®cally, recent developments in eral area cladograms.'' historical biogeography suggest to us that the degree It was evident immediately that in practice the of integration is something that changes in predictable search for topological congruence would be confronted ways through the ontogeny of a discipline, re¯ecting by ambiguity associated with widespread taxa, redun- shifts in the perceived reliability of potentially relevant dant distributions, and missing taxa (e.g., Nelson and information. Platnick, 1981; Page, 1990). This led to the develop- ment of several methods, especially ``component anal- The integration of phylogeny and biogeography ysis'' (e.g., Nelson and Platnick, 1981; Page, 1990, The fundamental idea underlying cladistic bioge- 1993a) and ``Brooks parsimony analysis'' (e.g., ography is that general patterns demand general ex- Brooks, 1981, 1985, 1990). Component analysis planations. Leon Croizat (e.g., 1958, 1964) champi- searches for general area cladograms under a variety oned this outlook in seeking generalized ``tracks'' link- of a priori assumptions about the evidential signi®- cance of widespread species, etc. Brooks parsimony analysis renders individual taxon area cladograms as a 1 From the Symposium on The Promise of Integrative Biology presented at the Annual Meeting of the Society for Integrative and matrix of binary characters and then combines these Comparative Biology, 2±6 January 2002, at Anaheim, California. in a single parsimony analysis. In general, deviations 2 E-mail: [email protected] from the emergent pattern of topological congruence 261 262 M. J. DONOGHUE AND BRIAN R. MOORE are attributed to lineage-speci®c processes, such as ex- tinction and dispersal (e.g., Brooks, 1990; Brooks and McLennan, 1991). The biogeographic problem is anal- ogous to the inference of other types of historical as- sociations, such as the coevolution of host-parasite systems and the ``gene tree/species tree'' problem, which have analogues to lineage-speci®c processes (e.g., ``dispersal'' has been equated with ``host switch- ing'' and ``horizontal transfer'': Page, 1993b, 1994; Maddison, 1995, 1997; Page and Charleston, 1998). More recently there have been a variety of meth- odological developments. From component analysis emerged ``reconciled tree analysis'' (Page, 1993b, 1994), ``three-item statements analysis'' (Nelson and Platnick, 1981; Nelson and Ladiges, 1991a, b), and FIG. 1. The test of time in historical biogeography. Congruence ``paralogy-free subtree analysis'' (Nelson and Ladiges, and incongruence refer to clades that diversi®ed synchronously and 1996; Ladiges, 1998). Brooks parsimony analysis has exhibit the same or different taxon area relationships, respectively. also been extended, especially in what has been called Pseudo-congruence and pseudo-incongruence refer to clades that di- ``secondary BPA'' (e.g., Brooks, 1990; Brooks and versi®ed at different times, but exhibit the same or different taxon area relationships, respectively (see text for discussion). McLennan, 1991; Brooks et al., 2001). The details of these methods have been reviewed elsewhere (e.g., Morrone and Carpenter, 1994; Mor- A(B,C). This phenomenon (e.g., Hafner and Nadler, rone and Crisci, 1995; Humphries and Parenti, 1999). 1990; Page, 1990, 1991; Hedges et al., 1994) has been We mention them here to emphasize that while a num- called ``pseudo-congruence'' (Cunningham and Col- ber of methods have been developed, they are funda- lins, 1994). The same area relationships were obtained, mentally similar with respect to the present arguments. but at different times, and presumably with different Most importantly, they are ``topology based,'' meaning underlying causes. that general patterns are sought by comparing or com- The other possible ®nding is that the topologies of bining only the topologies of component taxon area the area cladograms for the two groups are in con¯ict, cladograms. Although there has been discussion of the say, A(B,C) versus (A,B)C. This phenomenon (Page, ages of lineages, especially in the context of explaining 1990, 1991) may be referred to as ``pseudo-incongru- observed con¯icts among area cladograms, cladistic ence.'' The cladistic biogeographic interpretation biogeographic methods have not directly incorporated would be that the two groups were subjected to the information on the absolute timing of diversi®cation same geological events but somehow responded dif- in the component lineages. They also do not directly ferently. However, knowledge of the ages of the incorporate inferences on the place of origin or the groups indicates that they actually diversi®ed in re- directions of migration within particular lineages. sponse to different events. These concepts are illustrated in Figure 1. ``Con- WHAT'S WRONG WITH EXCLUDING TEMPORAL gruence'' and ``incongruence'' apply to lineages that INFORMATION? diversi®ed during the same time period and might Brie¯y, knowledge of the timing of events distin- therefore have been subjected to the same biogeo- guishes those groups that diversi®ed during the same graphic causes. Congruence suggests a common cause, time period, and therefore might have experienced the whereas incongruence implies lineage-speci®c phe- same causal events, from those that diversi®ed during nomena, such as dispersal and extinction. In contrast, different time periods and require different causal ex- ``pseudo-congruence'' and ``pseudo-incongruence'' re- planations (e.g., Grande, 1985; Hafner and Nadler, fer to cases in which lineages are not temporally co- 1990; Page, 1990, 1991, 1996; Cunningham and Col- incident. Under these circumstances one can only ob- lins, 1994; Hedges et al., 1994; Huelsenbeck et al., tain false-positive results or false-negative results. To
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