The voice of historical biogeography

Historical biogeography is going through an extraordinary revolution concerning its foundations, basic concepts, methods, and relationships to other disciplines of comparative biology. There are external and internal forces that are shaping the present of historical biogeography. The external forces are: global tectonics as the dominant paradigm in geosciences, cladistics as the basic language of comparative biology and the biologist's perception of biogeography. The internal forces are: the proliferation of competing articulations, recourse to philosophy and the debate over fundamentals. The importance of the geographical dimension of life's diversity to any understanding of the history of life on earth is emphasized. Three different kinds of processes that modify the geographical spatial arrangement of the organisms are identified: extinction, dispersal and vicariance. Reconstructing past biogeographic events can be done from three different perspectives: (1) the distribution of individual groups (taxon biogeography) (2) areas of endemism (area biogeography), and (3) biotas (spatial homology). There are at least nine basic historical biogeographic approaches: centre of origin and dispersal, panbiogeography, phylogenetic biogeography, cladistic biogeography, phylogeography, parsimony analysis of endemicity, event-based methods, ancestral areas, and experimental biogeography. These nine approaches contain at least 30 techniques (23 of them have been proposed in the last 14 years). The whole practice and philosophy of biogeography depend upon the development of a coherent and comprehensive conceptual framework for handling the distribution of organisms and events in space.     

Panbiogeography and conservation science in New Zealand

Abstract

New Zealand’s official conservation science framework, the Protected Natural Areas Programme (PNA), is evaluated in relation to current biogeographic/systematic methods and principles with reference to philosophy, taxonomy, information content, and historical geological/biological relations. The PNA Programme and its parent International Union for Conservation of Nature and Natural Resources (IUCN) framework is based on a phenetic system of classification that does not reflect process (phylogenetic) characters of ecological systems. The PNA philosophy of conservation science is shown to be essentialist, rooted in traditional medieval Western metaphysics, and out of step with current developments in biogeography, systematics, and ecology. Panbiogeography is an appropriate global perspective for developing a conservation science because it meets the requirements of homology, monophyly, increased information content, and empirical testability whereas the PNA programme does not. Establishment of a Panbiogeographic Track Atlas is proposed as a suitable conservation framework for historical ecology and biogeography. The atlas could provide an empirical natural resource inventory to identify priority areas for conservation at an economically acceptable cost compared to the PNA Programme. Application of cladistic techniques to ecological and biogeographic patterns in relation to the Atlas can provide aphylogenetically sound hierarchical classification for conservation science. The potential benefits of the panbiogeographic approach for conservation education and tourism are discussed.     

Historical Biogeography: Evolution in Time and Space

Biogeography is the discipline of biology that studies the present and past distribution patterns of biological diversity and their underlying environmental and historical causes. For most of its history, biogeography has been divided into proponents of vicariance explanations, who defend that distribution patterns can mainly be explained by geological, tectonic-isolating events; and dispersalists, who argue that current distribution patterns are largely the result of recent migration events. This paper provides an overview of the evolution of the discipline from methods focused on finding general patterns of distribution (cladistic biogeography), to those that integrate biogeographic processes (event-based biogeography), to modern probabilistic approaches (parametric biogeography). The latter allows incorporating into biogeographic inference estimates of the divergence time between lineages (usually based on DNA sequences) and external sources of evidence, such as information on past climate and geography, the organism fossil record, or its ecological tolerance. This has revolutionized the discipline, allowing it to escape the dispersal versus vicariance dilemma and to address a wider range of evolutionary questions, including the role of ecological and historical factors in the construction of biomes or the existence of contrasting patterns of range evolution in animals and plants.     

Paralogy and the Centre of Origin Concept

Ancestral area methodology, applied to finding the centre of origin, conflicts with most cladistic biogeographic methods since it uses, not reduces, paralogy. A new term, area cladistics, is herein proposed as an efficient paralogy-free (or reduced) method under the three-item philosophy that currently exists with other methods under the broad term cladistic biogeography.     

Toward an integrative historical biogeography

Cladistic biogeographic methods remain susceptible to the confoundingeffects of "pseudo-congruence" and "pseudo-incongruence" becausethey were not designed to incorporate information on the absolutetiming of the diversification of lineages. Consequently, resultsfrom cladistic biogeographic studies are difficult to interpretand cannot be confidently attributed to any particular cause.We illustrate these points with concrete examples, paying specialattention to recent work on the biogeography of the NorthernHemisphere, and outline ways in which topological and temporalinformation might be better integrated. The development of historicalbiogeography over the last few decades provides general insightsinto the nature of integration through the life of a discipline.     

When phylogenetics met biogeography: Willi Hennig,Lars Brundin and the roots of phylogenetic and cladistic biogeography

Willi Hennig's (Beitr. Ent. 1960, 10, 15) Die Dipteren-Fauna von Neuseeland als systematisches und tiergeographisches Problem applied a phylogenetic approach to examine the distributional patterns exhibited by the Diptera of New Zealand. Hennig showed how phylogenetic trees may be used to infer dispersal, based on the progression and deviation rules, and also discussed the existence of vicariance patterns. The most important author who applied Hennig's phylogenetic biogeography was Lars Brundin, when analysing the phylogenetic relationships of two taxa of Chironomidae (Diptera) and using them to examine the biogeographic relationships of Australia, New Zealand, South America and South Africa. The relevance of Brundin's contribution was noted by several authors, as it began the cladistic or vicariance approach to biogeography, that implies the discovery of vicariance events shared by different monophyletic groups. Both phylogenetic and cladistic biogeography have a place in contemporary biogeography, the former for analysing taxon biogeography and the latter when addressing Earth or biota biogeography. The recent use of the term “phylogenetic biogeography” to refer to a posteriori methods of cladistic biogeography is erroneous and should be avoided.     

LisBeth: New cladistics for phylogenetics and biogeography

Within phylogenetics, two methods are known to implement cladistics: parsimony or maximum parsimony (MP) and three-item analysis (3ia). Despite the lack of suitable software, 3ia is occasionally used in systematic, and more regularly, in historical biogeography. Here, we present LisBeth, the first and only phylogenetic/biogeographic program freely available that uses the 3ia approach and offer some insights into its theoretical propositions. LisBeth does not rely on the conventional taxon/character matrix. Instead, characters are represented as rooted trees. LisBeth performs 3ia analyses based on maximum congruence of three-item statements and calculates the intersection tree (which differs from usual consensus). In biogeography, it applies the transparent method to handle widespread taxa and implements paralogy-free subtree analysis to remove redundant distributions. For the sake of interoperability, LisBeth may import/export characters from/to matrix in NEXUS format, allowing comparison with other cladistic programs. LisBeth also imports phylogenetic characters from Xper² knowledge bases.     

历史生物地理学的进展

近年来历史生物地理学的进展主要是隔离分化学派的进展。对隔离分化学派的几个分支学派,特别是分支生物地理学和泛生物地理学在理论和方法方面的进展作了简要回顾和介绍。最近十年来分子手段广泛应用于历史生物地理学研究的各个方面,尤其是谱系生物地理学的快速崛起是历史生物地理学的一个明显特征。对本学科的发展做了初步的展望。     

Cladistic and phylogenetic biogeography: the art and the science of discovery

All methods used in historical biogeographical analysis aim to obtain resolved area cladograms that represent historical relationships among areas in which monophyletic groups of taxa are distributed. When neither widespread nor sympatric taxa are present in the distribution of a monophyletic group, all methods obtain the same resolved area cladogram that conforms to a simple vicariance scenario. In most cases, however, the distribution of monophyletic groups of taxa is not that simple. A priori and a posteriori methods of historical biogeography differ in the way in which they deal with widespread and sympatric taxa. A posteriori methods are empirically superior to a priori methods, as they provide a more parsimonious accounting of the input data, do not eliminate or modify input data, and do not suffer from internal inconsistencies in implementation. When factual errors are corrected, the exemplar presented by M.C. Ebach & C.J. Humphries (Journal of Biogeography, 2002, 29 , 427) purporting to show inconsistencies in implementation by a posteriori methods actually corroborates the opposite. The rationale for preferring a priori methods thus corresponds to ontological rather than to epistemological considerations. We herein identify two different research programmes, cladistic biogeography (associated with a priori methods) and phylogenetic biogeography (associated with a posteriori methods). The aim of cladistic biogeography is to fit all elements of all taxon–area cladograms to a single set of area relationships, maintaining historical singularity of areas. The aim of phylogenetic biogeography is to document, most parsimoniously, the geographical context of speciation events. The recent contribution by M.C. Ebach & C.J. Humphries (Journal of Biogeography, 2002, 29 , 427) makes it clear that cladistic biogeography using a priori methods is an inductivist/verificationist research programme, whereas phylogenetic biogeography is hypothetico‐deductivist/falsificationist. Cladistic biogeography can become hypothetic‐deductive by using a posteriori methods of analysis.     

Critique of parsimony analysis of endemicity as a method of historical biogeography

Aim Assess the value of parsimony analysis of endemism as either an a priori (cladistic) and an a posteriori (phylogenetic) method of historical biogeography. Location World‐wide. Methods Parsimony analysis of endemicity (PAE) and Brooks parsimony analysis (BPA). Results Parsimony analysis of endemicity is capable of finding correct and unambiguous area relationships only under scenarios of vicariance in combination with non‐response to vicariance or extinction. An empirical comparison between PAE and BPA, using the poeciliid fish genera Heterandria and Xiphophorus, demonstrates that PAE fails to document much of the historical complexity in this relatively simple system. Main conclusions The a priori assumptions of PAE are far more restrictive than those made by other a priori methods, limiting its utility as a method of cladistic biogeography. The inability of PAE to detect perfect vicariance or biogeographical histories involving dispersal, renders it unsuitable as a method of phylogenetic biogeography.     

历史生物地理学进展

生物地理学研究动植物的地理分布。历史生物地理学重建生物区系历史。分替理论的复兴动摇了散布理论的上百年统治。最近10年主要是分替理论推动了历史生物地理学,出现了多个途径——种系发生物地理学、分支分替生物地理学、特有性的俭吝分析和泛生物地理学。岛屿生物地理学理论有了改进和严格的实验检验;庇所学说产生了新的模型。最后就我国如何发展生物地理学提出了对策措施。     

Fossil biogeography: a new model to infer dispersal,extinction and sampling from palaeontological data

Methods in historical biogeography have revolutionized our ability to infer the evolution of ancestral geographical ranges from phylogenies of extant taxa, the rates of dispersals, and biotic connectivity among areas. However, extant taxa are likely to provide limited and potentially biased information about past biogeographic processes, due to extinction, asymmetrical dispersals and variable connectivity among areas. Fossil data hold considerable information about past distribution of lineages, but suffer from largely incomplete sampling. Here we present a new dispersal–extinction–sampling (DES) model, which estimates biogeographic parameters using fossil occurrences instead of phylogenetic trees. The model estimates dispersal and extinction rates while explicitly accounting for the incompleteness of the fossil record. Rates can vary between areas and through time, thus providing the opportunity to assess complex scenarios of biogeographic evolution. We implement the DES model in a Bayesian framework and demonstrate through simulations that it can accurately infer all the relevant parameters. We demonstrate the use of our model by analysing the Cenozoic fossil record of land plants and inferring dispersal and extinction rates across Eurasia and North America. Our results show that biogeographic range evolution is not a time-homogeneous process, as assumed in most phylogenetic analyses, but varies through time and between areas. In our empirical assessment, this is shown by the striking predominance of plant dispersals from Eurasia into North America during the Eocene climatic cooling, followed by a shift in the opposite direction, and finally, a balance in biotic interchange since the middle Miocene. We conclude by discussing the potential of fossil-based analyses to test biogeographic hypotheses and improve phylogenetic methods in historical biogeography.     

替代学派生物地理学几种研究方法简介

历史生物地理学就其基本原理、概念和研究方法可大致分为以过程为导向的传播学派和以式样为导向的替代学派。本文简要介绍替代学派的一些基本原理和概念,对最近半个世纪以来发展的几种替代学派生物地理学方法：泛生物地理学、更新世森林避难所理论、分支生物地理学以及特有性的简约性分析法的原理、方法作了简要的介绍和评论。     

Vagility: The Neglected Component in Historical Biogeography

The conceptual gap between ecological and historical biogeography is wide, although both disciplines are concerned with explaining how distributions have been shaped. A central aim of modern historical biogeography is to use a phylogenetic framework to reconstruct the geographic history of a group in terms of dispersals and vicariant events, and a number of analytical methods have been developed to do so. To date the most popular analytical methods in historical biogeography have been parsimony-based. Such methods can be classified into two groups based on the assumptions used. The first group assumes that vicariance between two areas creates common patterns of disjunct distributions across several taxa whereas dispersals and extinctions generate clade specific patterns. The second group of methods assumes that passive vicariance and within-area speciation have a higher probability of occurrence than active dispersal events and extinction. Typically, none of these methods takes into account the ecology of the taxa in question. I discuss why these methods can be potentially misleading if the ecology of the taxon is ignored. In particular, the vagility or dispersal ability of taxa plays a pivotal role in shaping the distributions and modes of speciation. I argue that the vagility of taxa should be explicitly incorporated in biogeographic analyses. Likelihood-based methods with models in which more realistic probabilities of dispersal and modes of speciation can be specified are arguably the way ahead. Although objective quantification will pose a challenge, the complete ignorance of this vital aspect, as has been done in many historical biogeographic analyses, can be dangerous. I use worked examples to show a simple way of utilizing such information, but better methods need to be developed to more effectively use ecological knowledge in historical biogeography.     

Titi monkey biogeography: Parallel Pleistocene spread by Plecturocebus and Cheracebus into a post‐Pebas Western Amazon

Titi monkeys, subfamily Callicebinae, are a diverse, species‐rich group of Neotropical primates with an extensive range across South America. Their distribution in space and time makes them an interesting primate model for addressing questions of Neotropical historical biogeography. Our aim was to reconstruct the biogeographic history of Callicebinae to better understand their diversification patterns and the history of their colonisation of South America since the late Miocene. We reconstructed a time‐calibrated phylogeny of 19 titi species under Bayesian inference using two mitochondrial and 11 nuclear loci. Species were assigned across eight Neotropical areas of endemism, and statistical biogeographic methods implemented in BioGeoBEARS were employed to estimate ancestral areas using 12 biogeographic models. Our results indicate that the most recent common ancestor to extant titi monkeys was widespread from the present‐day Andean foothills in the Colombian Amazon, through the wet and dry savannas of Bolivia and Brazil, to the southern Atlantic forest of eastern Brazil. Genus‐level divergences were characterised by vicariance of ancestral range in the late Miocene. Species‐level diversification in Cheracebus and the Plecturocebus moloch group occurred as they spread across the Amazon in the Pleistocene and were largely characterised by a sequential, long‐distance “island‐hopping” dispersal model of speciation from a narrow area of origin through jump dispersal across rivers. This study comprises the first large‐scale investigation of the evolutionary history of titi monkeys in the context of Amazonian and South American historical biogeography and sheds light on the processes that generated the great diversity found among Callicebinae.     

Review of plant biogeographic studies in Brazil

Molecular phylogenetic studies have become a major area of interest in plant systematics, and their impacts on historical biogeographic hypotheses are not to be disregarded. In Brazil, most historical biogeographic studies have relied on animal phylogenies, whereas plant biogeographic studies have largely lacked a phylogenetic component, having a limited utility for historical biogeography. That country, however, is of great importance for most biogeographic studies of lowland tropical South America, and it includes areas from a number of biogeographic regions of the continent. Important biogeographic reports have been published as part of phylogenetic studies, taxonomic monographs, and regional accounts for small areas or phytogeographic domains, but the available information is subsequently scattered and sometimes hard to find. In this paper we review some relevant angiosperm biogeographic studies in Brazil. Initially we briefly discuss the importance of other continents as source areas for the South American flora. Then we present a subdivision of Brazil into phytogeographic domains, and we cite studies that have explored the detection of biogeographic units （areas of endemism） and how they are historically related among those domains. Examples of plant taxa that could be used to test some biogeographic hypotheses are provided throughout, as well as taxa that exemplify several patterns of endemism and disjunction in the Brazilian angiosperm flora.     

The reform of palaeontology and the rise of biogeography – 25 years after 'ontogeny, phylogeny, paleontology and the biogenetic law' (Nelson, 1978)

Aim To document the historical development of cladistics and the roles palaeontology and biogeography played in establishing coherent concepts of phylogenetic relationships focusing on some aspects of the contributions of Gareth Nelson. Conclusions Nelson's reformulation of the threefold parallelism provides a rationale for investigating phylogeny, replacing the central role palaeontology once played with biogeography, adding a spatial dimension to the concept of phylogeny. This approach to phylogeny replaces the old ‘transformationist’ view with the cladistic view, the latter dependent on discovering relationships among taxa. Numerical phylogenetic methods are inherently ‘transformationist’ and have replaced stratigraphy as the key to phylogenetic relationships. Numerical methods in systematics and biogeography are inherently transformational and suffer the same problems as the old palaeontology.     

New Guinea: A Correlation between Accreting Areas and Dispersing Sapindaceae

A correlation between accreting (hybridizing) areas and dispersing taxa (several genera of Southeast Asian/Australian Sapindaceae) is theoretically impossible in cladistic biogeography. However, in particular circumstances (primitive absence followed by colonization and speciation) cladistic methods can reconstruct (part of) the historical sequence of accretion. In this example, the phases in the accretion history of more than 30 terranes of the northern half of New Guinea correspond reasonably well with the generalized area cladogram of the Sapindaceae.     

Panbiogeography from tracks to ocean basins: evolving perspectives

Misconceptions arising from efforts to translate panbiogeography into terms used in other biogeographic and evolutionary theories are discussed with respect to 5 ) critique of panbiogeography. Croizat’s rejection of ‘Darwinian dispersal’ applies only to efforts to utilize this concept as a general explanation for biogeographic patterns. The conceptual difference between distribution and panbiogeographic dispersal maps is illustrated to show that Croizat did not synonymize distribution and dispersal. Croizat’s position on continental drift and plate tectonics does not support 5 claim that Croizat ‘for a long time’ refused to accept the theory of plate tectonics. The methodological relationship between panbiogeographic analysis and geology suggests an independence of methodology that prevents geological theory from falsifying panbiogeographic predictions. Panbiogeographic predictions for the eastern Pacific are shown to be in agreement with current historical geological models. Claims by 5 that the panbiogeographic method is variable and questionable are evaluated with respect to the biogeographic homology of primitive frogs, ratite birds, and southern beeches to demonstrate the consistent application of minimal distance, main massing, phylogenetic affinity and baseline criteria. Panbiogeographic classification concepts are contrasted with the Darwinian system (supported by Cox) utilizing a concept of unitary geographical area based on the language of Roman military rule. Inconsistent positions expressed in recent critiques of panbiogeography may indicate an underlying and implicit acceptance of the empirical and theoretical progress generated by panbiogeography within modern biogeography. ‘The formation of groups has an invigorating effect in all spheres of human striving, perhaps mostly due to the struggle between the convictions and aims represented by the different groups’ (Einstein, 1938. Collier’s, 26 November).     

The dynamics of biogeographic ranges in the deep sea

Anthropogenic disturbances such as fishing, mining, oil drilling, bioprospecting, warming, and acidification in the deep sea are increasing, yet generalities about deep-sea biogeography remain elusive. Owing to the lack of perceived environmental variability and geographical barriers, ranges of deep-sea species were traditionally assumed to be exceedingly large. In contrast, seamount and chemosynthetic habitats with reported high endemicity challenge the broad applicability of a single biogeographic paradigm for the deep sea. New research benefiting from higher resolution sampling, molecular methods and public databases can now more rigorously examine dispersal distances and species ranges on the vast ocean floor. Here, we explore the major outstanding questions in deep-sea biogeography. Based on current evidence, many taxa appear broadly distributed across the deep sea, a pattern replicated in both the abyssal plains and specialized environments such as hydrothermal vents. Cold waters may slow larval metabolism and development augmenting the great intrinsic ability for dispersal among many deep-sea species. Currents, environmental shifts, and topography can prove to be dispersal barriers but are often semipermeable. Evidence of historical events such as points of faunal origin and climatic fluctuations are also evident in contemporary biogeographic ranges. Continued synthetic analysis, database construction, theoretical advancement and field sampling will be required to further refine hypotheses regarding deep-sea biogeography.