Two Requirements for Obtaining Valid Common Patterns under Different Assumptions in Vicariance Biogeography

In vicariance biogeography, widespread or sympatric taxa can be dealt with under assumptions 0, 1, and 2. Data from cladogenetic relationships among taxa of a monophyletic group and their distribution over areas are assumed, in the order 0 → 1 → 2, to represent decreasing information about vicariance events. A less strict assumption carries a larger solution set, i.e., the number of possible area cladograms increases with the decrease in strictness of the assumption applied. We formulate two requirements for obtaining valid general area cladograms from data of several monophyletic groups of taxa. First, the assumptions, and with them the sets of area cladograms derived under these assumptions, should be inclusive. Second, sets of single group area cladograms should be compared for different monophyletic groups under a single assumption. When these two requirements are met, area cladograms become consistent with respect to the processes (vicariance, extinction, and dispersal) that are a priori assumed. The explanatory power increases for any particular monophyletic group of taxa when the set of valid general area cladograms contains a subset of area cladograms derived under a more strict assumption. We discuss examples from literature of how violation of these two requirements affects the results.     

Methods in Vicariance Biogeography: Assessment of the Implementations of Assumptions 0, 1, and 2

As we have argued previously, for the valid derivation of general area cladograms in vicariance biogeography, two requirements should be met. First, sets of area cladograms derived under assumptions 0, 1 and 2 should be inclusive (requirement I). Second, general area cladograms should be based on area cladograms, for different monophyletic groups, derived under the same assumption (requirement II). We now assess for their actual implementation of assumptions A0, A1, and A2 and for the extent to which they meet requirements I and II, the following methods (and correlated computer programs): Component Compatibility Analysis (CAFCA), Brooks Parsimony Analysis (PAUP), Component Analysis (Component 1.5), Reconciled Tree Analysis (Component 2.0), and Three Area Statement Analysis (TAS). For this purpose we use empirical ( Heterandria, Xiphophorus, Cyttaria, Eriococcus/Madarococcus ) and theoretical data sets. All programs appear to violate, to a different degree, requirement I (deriving inclusive sets of area cladograms under assumptions) when dealing with sympatric taxa under A1 or A2. Dealing with sympatric taxa a posteriori only prevents this violation. All programs examined appear to meet requirement II (deriving general area cladograms under a single assumption).     

Measures for Obtaining Inclusive Sets of Area Cladograms under Assumptions Zero, 1, and 2 with Different Methods for Vicariance Biogeography

We present modifications to computer programs of a posteriori and a priori methods for vicariance biogeography that enable them to obtain inclusive sets of area cladograms under assumptions zero, 1, and 2. With CAFCA (Component Compatibility Analysis) an upper limit for area cladogram selection by the number of steps is not always sufficient for inclusive sets. CAFCA needs additionally a lower limit for the number of components used to derive area cladograms when noninclusion arises because CAFCA selects area cladograms with different resolutions. PAUP (Brooks Parsimony Analysis) derives inclusive sets when it selects area cladograms under assumptions zero, 1, and 2 by using an upper limit for the number of steps and not collapsing unsupported nodes. For the computer programs Component 1.5 (Component Analysis), Component 2.0 (Reconciled Tree Analysis), and TAS (Three Area Statement Analysis) we suggest a two-step procedure for inclusive sets. The first step involves dealing with widespread taxa a priori under assumptions zero, 1, or 2. The second step involves dealing with sympatric taxa "as is" (Component 1.5 and TAS) or by tree reconciliation using an upper limit for the number of losses (Component 2.0).     

PACT: an efficient and powerful algorithm for generating area cladograms

Aim To introduce and describe the functioning of a new algorithm, phylogenetic analysis for comparing trees (PACT), for generating area cladograms that provide accurate representation of information contained in taxon–area cladograms. Methods PACT operates in the following steps. Convert all phylogenies to taxon–area cladograms. Convert all taxon–area cladograms to Venn diagrams. Choose any taxon–area cladogram from the set of taxon–area cladograms to be analysed and determine its elements. This will be the template area cladogram. Select a second taxon–area cladogram. Determine its elements. Document which elements in the second tree occur in the template tree (denoted by ‘Y’) and which do not (denoted by ‘N’). Each ‘Y’ indicates a match with previous pattern and these are combined. Each ‘N’ is a new element and is attached to the template area cladogram at the node where it is linked with a Y. This requires two rules: (1) ‘Y + Y = Y’ (combine common elements) as long as they are connected at the same node; and (2) ‘Y + N = YN’ (add novel elements to the template area cladogram at the node where they first appear). Once the novel elements in the second taxon–area cladogram have been added to the template area cladogram, see if any of them can be further combined. This requires three additional rules: (1) ‘Y(Y? = Y(Y?’ (do not combine Y's if they are attached at different nodes on the template area cladogram); (2) ‘Y + YN = YN’ (Y is part of group YN); and (3) ‘YN + YN = YNN’ (Y is the same for each, but each N is different). Repeat for all available taxon–area cladograms. Results Three exemplars demonstrate that PACT provides the most accurate area cladograms for vicariance‐driven biotic diversification, dispersal‐driven biotic diversification and taxon pulse‐driven biotic diversification. PACT can also be used as an a priori method of biogeographical analysis. Main conclusions PACT embodies all the strong points and none of the weaknesses of previously proposed methods of historical biogeography. It is most useful as an a posteriori method, but it is also superior to all previous a priori methods because it does not specify costs, or weights or probabilities, or likelihoods of particular biogeographical processes a priori and is thus sensitive to clade‐specific historical contingencies.     

ON CONSENSUS, COLLAPSIBILITY, AND CLADE CONCORDANCE

Abstract — Consensus in cladistics is reviewed. Consensus trees, which summarize the agreement in grouping among a set of cladograms, are distinguished from compromise trees, which may contain groups that do not appear in all the cladograms being compared. Only a strict or Nelson tree is an actual consensus. This distinction has implications for the concept of support for cladograms: only those branches supported under all possible optimizations are unambiguously supported. We refer to such cladograms as strictly supported, in contrast to the semistrictly (ambiguously) supported cladograms output by various current microcomputer programs for cladistic analysis. Such semistrictly supported cladograms may be collapsed, however, by a variety of options in various programs. Consideration of collapsibility and optimization on multifurcations leads to some conclusions on the use of consensus. Consensus tree length provides information about character conflict that occurs between, not within, cladograms. We propose the clade concordance index, which employs the consensus tree length to measure inter-cladogram character conflict for all characters among a set of cladograms.     

Parsimony analysis of endemicity (PAE) of Mexican terrestrial mammals at different area units: when size matters

Abstract Aim Parsimony analysis of endemicity (PAE) is a biogeographical method that uses a parsimony algorithm to obtain an area cladogram, based on taxa inhabiting the study areas. We compare its performance at different geographical units (½° and 1° quadrats, ecoregions and biogeographical provinces) to analyse distributional patterns of Mexican terrestrial mammals, in order to assess the importance of the size of area units. Location The area analysed corresponds to Mexico. Methods Parsimony analyses were based on 56,859 collection records, corresponding to 703 genera, species and subspecies. Four data matrices were constructed for: (1) 716 quadrats of ½° latitude × ½° longitude, (2) 230 quadrats of 1° latitude × 1° longitude, (3) forty‐five ecoregions and (4) fourteen biogeographical provinces. Results For the ½° quadrat matrix, we obtained six cladograms of 17,138 steps. For the 1° quadrat matrix, we obtained five cladograms (strict consensus with 9394 steps). For the matrix of ecoregions, we obtained twelve cladograms (strict consensus cladogram with 3009 steps). For the provinces, we obtained a single cladogram with 1603 steps. Main conclusions The best results were obtained with natural areas instead of quadrats. There seems to exist a trend to decrease the absolute number of steps and an increase in the absolute and relative number of synapomorphies as the size of the area units decreases, although this does not necessarily occur for the number of cladograms.     

Marine Area Relationships from Twenty Sponge Phylogenies. A Comparison of Methods and Coding Strategies

Published phylogenies of 20 marine sponge groups are used to build general area cladograms of marine areas of endemism under three different methods for which algorithms adapted for personal computers are available, viz. COMPONENT, BPA and TAS, and two different coding strategies, Assumption 0 (A0) and "no assumption" (NA). The latter is a recently proposed procedure for handling the distributions of widespread taxa by treating these as separate areas of endemism, rather than as suites of smaller constituent areas. The 20 phylogenies contained large numbers of problem data which prevented an exhaustive search for all possible equally "best" general area cladograms. The Nelson consensus trees and their equivalents in parsimony analysis for all six attempts (viz. three different methodologies under two different coding strategies) were compared using their fit with the 20 sponge phylogenies as a measure of quality. Fit was determined using the number of "cospeciations" between a general area cladogram and a taxon area cladogram computed with TreeMap 1.0. No single method or coding strategy yielded a clearly better fit, each cladogram fitting variously better or worse with various phylogenies. In general, fit with NA coding was higher than with A0 coding, but random tree tests failed to generate statistically significant support for the conclusion that NA coding improves fit. Assuming that available sponge phylogenies are representative of marine benthic groups, software and hardware limitations are serious obstacles to a successful development of marine general area cladograms under any method or coding strategy.     

Intraspecific phylogeography of the slender madtom: the complex evolutionary history of the Central Highlands of the United States

A number of different biogeographical studies of the Central Highlands of the United States have yielded conflicting area cladograms. We estimate the mtDNA phylogeny of populations of the slender madtom, Noturus exilis, a clear-water stream catfish. The goal is to compare population relationships to those reported in previous studies that used upland, stream-dwelling vertebrates. A region of the NADH dehydrogenase subunit 4 gene, along with adjacent tRNAs, was sequenced for population samples from 21 different Central Highlands rivers. Sequence difference among 39 haplotypes ranged from 0.1% to 4.8%. Most haplotypes were restricted to specific rivers and mapped well onto geography. Slender madtoms from different drainages contained mostly monophyletic groups of haplotypes genetically divergent from haplotypes found in other drainages, although a few haplotypes were found in well-separated drainages. The area cladogram for the slender madtom was not similar to any of the other cladograms for other species and species groups from the area. We discuss a variety of methodological and biological reasons for the discordance, and suggest that some of the discrepancies may be resolved by the sequencing of multiple genes per species. We recommend that more, and more extensive, intraspecific phylogeography studies should be conducted for species living in the Central Highlands rivers.     

The unfalsifiability of cladograms and its consequences

Popper's falsificationism provides the normative reference system in recent discussions regarding theory and methodology of systematics. According to Popper, the falsifiability of a hypothesis represents a necessary precondition for its corroborability. It is shown that cladograms, independent of “strict”, “methodological” or “sophisticated” falsification, are not falsifiable in principle. No present observation is prohibited by any tree hypothesis and, thus, no Popperian test of cladograms exists. It is shown that the congruence test, which is commonly said to represent a Popperian test of cladograms, instead tests sets of apomorphy hypotheses. Three different strategies that have been proposed to circumvent this problem are discussed and refuted: (1) referring to Popper's convention to renounce ad hoc maneuvers; (2) referring to Popper's treatment of probability hypotheses; and (3) decoupling corroboration from falsification. As a consequence, within a Popperian framework the unfalsifiability of cladograms implies that cladograms cannot explain any present day observation and, thus, represent metaphysical hypotheses. However, Popper's falsificationism has been criticized and questioned by many philosophers before and it seems to be about time that phylogeneticists develop their own philosophy of phylogenetics that meets their specific requirements of a historical science that is not seeking for universal laws and regularities, but instead reconstructing particular historical events. © The Willi Hennig Society 2007.     

A PRELIMINARY PHYLOGENETIC AND BIOGEOGRAPHIC ANALYSIS OF THE DYAKIIDAE (GASTROPODA: STYLOMMATOPHORA) AND A BIOGEOGRAPHIC ANALYSIS OF OTHER SUNDALAND TAXA

Abstract — The phylogeny of the Dyakiidae, an early diverging lineage of the Limacoidea sensu lalo , is reconstructed. In the Dyakiidae a unique transformation series of different genital accessory organs, the so-called "stimulators", is preserved which is an important argument for the hypothesis that the different stimulator types of the Stylommatophora are homologous. The biogeographic distribution patterns of the genera of the Dyakiidae are summarized and the ancestral areas of the major clades are analysed. The analysis of the area cladograms of the Dyakiidae and of several butterfly and heteroptera genera occurring in Sundaland revealed neither a general pattern of relationship between the areas of endemism in Sundaland, nor a general pattern of relationship between Sundaland and other areas. The area cladogram of the rather immobile Dyakiidae endemic to Sundaland might reflect older events than the area cladograms of the more mobile and widespread butterfly and plant bug genera. The general incompatibility of the area cladograms involving Sundaland taxa suggests that dispersal across barriers has played a major role in the historical biogeography of the analysed groups and challenges the hypothesis of vicariance biogeography that the distribution patterns of organisms are largely due to the fragmentation of an ancestral biota.     

WIDESPREAD TAXA, SYMPATRY, DISPERSAL, AND AN ALGORITHM FOR RESOLVED AREA CLADOGRAMS

Abstract — It is argued that in area biogeographical studies, the information content of widespread taxa is comparable to that of endemic taxa; widespread taxa should therefore be treated under Assumption 0. For areas of sympatry (redundant distributions) on the other hand, Assumption 2 is warranted because dispersal must always be invoked to explain sympatry. It is argued that all occurrences in an area of sympatry should be treated equally, regardless of whether the taxa are widespread or endemic, and regardless of whether the taxa occupy terminal or basal positions on the cladogram. An algorithm for translation of taxon-area cladograms into resolved area cladograms under these conditions is given. As an example, this algorithm is applied to inocelliid snakeflies and gerrine pondskaters.     

A Posteriori and a Priori Methodologies for Testing Hypotheses of Causal Processes in Vicariance Biogeography

Methods used in vicariance biogeography fall into the categories of a posteriori methods (e.g., Component Compatibility Analysis and Brooks Parsimony Analysis) and a priori methods (e.g., Component Analysis, Reconciled Tree Analysis, and Three Area Statement Analysis). Each category corresponds to a particular methodology that arrives at general area cladograms by testing null hypotheses in a particular way. A posteriori methods assume the process of vicariance only (A0) as a common cause of the distribution of different monophyletic groups of taxa under the null hypothesis. Whenever a parsimony analysis of combined data from these monophyletic groups results in a general area cladogram with homoplasy, the null hypothesis is rejected and extinction and dispersal are invoked a posteriori as ad hoc process explanations. A priori methods assume not only vicariance (A0) but also combinations of vicariance with the processes of extinction (A1) and dispersal (A2) as possible causes of the distribution of the taxa of different monophyletic groups. Each assumed set of processes corresponds to a different null hypothesis. Under the assumption of independence and thus additivity of the processes involved, the sets of area cladograms obtained under A0, A1, and A2 from data of each monophyletic group must be inclusive (requirement I). Whenever no congruent area cladograms are found in the intersection of sets of area cladograms derived under the same assumption for different monophyletic groups (II), the corresponding null hypothesis is rejected.     

IN SEARCH OF A METHOD FOR CLADISTIC BIOGEOGRAPHY: AN EMPIRICAL COMPARISON OF COMPONENT ANALYSIS, BROOKS PARSIMONY ANALYSIS, AND THREE-AREA STATEMENTS

Abstract— Three quantitative cladistic biogeographic methods, namely, component analysis, Brooks parsimony analysis (BPA), and three-area statements (TAS) have been proposed for obtaining general area cladograms from taxon-area cladograms. Available programs implementing these methods include COMPONENT versions 1.5 and 2.0 for component analysis, TAS for three-area statements, and Hennig86 for analysing matrices for both three-area statements and BPA. Ten different data sets were analysed with these programs and items of error were used to evaluate the general area cladograms obtained. None of the computer implementations of the methods compared proved to be more effective than the others.
"…all methodologies, even the most obvious ones, have their limits…"
Feyerabend (1993:231)     

Widespread Taxa and Component 2.0

Resolved area cladograms found by heuristic search with COMPONENT 2.0, with the option “map widespread associates” turned off, correspond to Assumption 1 only if the widespread taxon occupies a terminal position on the taxon-area-cladogram. If the widespread taxon occupies a basal position, the resolved area cladograms correspond to Assumption 2.     

PHYLOGENY OF THE NEMERTEAN SUBCLASS PALAEONEMERTEA (ANOPLA, NEMERTEA)

Abstract— A cladistic analysis based on 50 morphological characters was performed for 49 of the 98 species currently assigned to the subclass Palaeonemertea (phylum Nemertea), and six additional undescribed species. Thirty-five species were excluded from the parsimony analysis because of the high number of unknowns in the character matrix, and one species since it was considered a nomen nudum . An initial analysis suggested that the subclass Hoplonemertea is the sistergroup to the clade Palaeo- and Heteronemertea and the ingroup cladograms are rooted using a paraphyletic outgroup based on this information. Seventy-two equally most parsimonious cladograms were found; the consistency index was low but tree-length distribution for the character set is skewed to the left, and the cladograms are invariably shorter than trees based on random data. These cladograms suggested a character transformation series for the cerebral organ where this complex character reappeared several times after being absent. We considered this biologically implausible and the final discussion is based on three cladograms, one step longer than the most parsimonious, where the evolution of this character appears to be more realistic. The cladistic analysis indicates that many previously recognized genera (e.g. Cephalothrix, Procephalothrix and Cephalotrichella ), and higher taxa, are paraphyletic. It furthermore indicates that the previously suggested hypothesis of the Archinemertea as a monophyletic sistertaxon to Palaeonemertea is unsupported.     

A Cladistic Analysis of Phenotypic Associations with Haplotypes Inferred from Restriction Endonuclease Mapping and DNA Sequence Data. III. Cladogram Estimation

We previously developed a cladistic approach to identify subsets of haplotypes defined by restriction endonuclease mapping or DNA sequencing that are associated with significant phenotypic deviations. Our approach was limited to segments of DNA in which little recombination occurs. In such cases, a cladogram can be constructed from the restriction site or sequence data that represents the evolutionary steps that interrelate the observed haplotypes. The cladogram is used to define a nested statistical design to identify mutational steps associated with significant phenotypic deviations. The central assumption behind this strategy is that any undetected mutation causing a phenotypic effect is embedded within the same evolutionary history that is represented by the cladogram. The power of this approach depends upon the confidence one has in the particular cladogram used to draw inferences. In this paper, we present a strategy for estimating the set of cladograms that are consistent with a particular sample of either restriction site or nucleotide sequence data and that includes the possibility of recombination. We first evaluate the limits of parsimony in constructing cladograms. Once these limits have been determined, we construct the set of parsimonious and nonparsimonious cladograms that is consistent with these limits. Our estimation procedure also identifies haplotypes that are candidates for being products of recombination. If recombination is extensive, our algorithm subdivides the DNA region into two or more subsections, each having little or no internal recombination. We apply this estimation procedure to three data sets to illustrate varying degrees of cladogram ambiguity and recombination.     

Cladistic biogeography of the moth tribe Cidariini (Lepidoptera, Geometridae) in the Holarctic and Indo-Chinese regions

A cladistic biogeographic analysis for the Holarctic and Indo-Chinese regions was undertaken based on seven genera of the tribe Cidariini: Cidaria Treitschke, Thera Stephens, Pennithera Viidalepp, Heterothera Inoue, Callabraxas Butler, Gandaritis Moore and Eulithis Httbner. Smallest coincident ranges of two species recognized 11 endemic areas. The study has two aims: to construct a hierarchical structure of those areas, and to recognize dispersal events. Under two assumptions [widespread taxa mapped (identical as assumption 0) and widespread taxa not mapped (identical as assumption 1)] the 11 endemic areas were mapped with 72 taxa. The best resolved area cladograms under the two assumptions differ in the placement of one endemic area, northern Europe. Area relationships found in this present analysis are congruent with the current landmass configurations: (North America, (Europe, (northern India, (southwestern Asia, (Baikal area, (south China, (Taiwan, (Russian Far East, Japan)))))))). These area cladograms postulate at least three vicariance events: (1) between North America and the Palaearctic; (2) western-eastern Palaearctic; (3) northern India–the rest of Asia. The approach to recognize dispersed taxa by pruning each taxon suggests that most dispersal events occurred in East Asia: from the Baikal area or south China to the Russian Far East; and from the Russian Far East to Japan. Relationships among endemic areas are briefly discussed.     

Application of parsimony analysis of endemicity in Amazonian biogeography: an example with primates

The distributions of 51 non-human primate species are used for Parsimony Analysis of Endemicity (PAE) to determine the relationships among 14 interfluvial regions in the Amazon basin, South America. Two most parsimonious cladograms were found. The strict consensus tree of these cladograms suggests an early separation between Lower Amazonia (eastern) and Upper Amazonia (western). The major clusters of interfluvial regions identified in the PAE cladogram are congruent with the areas of endemism delimited for birds. When interfluvial regions are converted into avian areas of endemism, the PAE cladogram is congruent with one of the two general areas cladograms suggested for Amazonia based on phylogenies of several clades of forest birds. Our analysis suggests that PAE can be used as a tool to objectively identify areas of endemism at an intra-continental scale as well as to make historical inferences. However, the value of a PAE cladogram in this latter application should be always evaluated by congruence with area cladograms built upon cladistic biogeography procedures.     

A Hypothesis-Independent Method of Character Weighting for Cladistic Analysis

Abstract— A hypothesis-independent method of weighting cladistir characters, based on character compatibility, is proposed. The method is used in two fashions, to generate cladograms, and to select from multiple minimum length cladograms. To illustrate the techniques, published data on Hoplinema (Coleoptera) are reanalysed. The method is contrasted with other weighting techniques which arc generally found to be hypothesis dependent.