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Here, we propose, prove mathematically and discuss maximum and minimum measures of maximum parsimony evolution across 12 discrete phylogenetic character types, classified across 4467 morphological and molecular datasets. Covered character types are: constant, binary symmetric, multistate unordered (non‐additive) symmetric, multistate linear ordered symmetric, multistate non‐linear ordered symmetric, binary irreversible, multistate irreversible, binary Dollo, multistate Dollo, multistate custom symmetric, binary custom asymmetric and multistate custom asymmetric characters. We summarize published solutions and provide and prove a range of new formulae for the algebraic calculation of minimum (m), maximum (g) and maximum possible (gmax) character cost for applicable character types. Algorithms for exhaustive calculation of m, g and gmax applicable to all classified character types (within computational limits on the numbers of taxa and states) are also provided. The general algorithmic solution for minimum steps (m) is identical to a minimum spanning tree on the state graph or minimum weight spanning arborescence on the state digraph. Algorithmic solutions for character g and gmax are based on matrix mathematics equivalent to optimization on the star tree, respectively for given state frequencies and all possible state frequencies meeting specified numbers of taxa and states. We show that maximizing possible cost (gmax) with given transition costs can be equivalent to maximizing, across all possible state frequency combinations, the lowest implied cost of state transitions if any one state is ancestral on the star tree, via the solution of systems of linear equations. The methods we present, implemented in the Claddis R package, extend to a comprehensive range, the fundamental character types for which homoplasy may be measured under parsimony using m, g and gmax, including extra cost (h), consistency index (ci), retention index (ri) or indices based thereon. [ABSTRACT FROM AUTHOR]

Although simulations have shown that implied weighting (IW) outperforms equal weighting (EW) in phylogenetic parsimony analyses, weighting against homoplasy lacks extensive usage in palaeontology. Iterative modifications of several phylogenetic matrices in the last decades resulted in extensive genealogies of datasets that allow the evaluation of differences in the stability of results for alternative character weighting methods directly on empirical data. Each generation was compared against the most recent generation in each genealogy because it is assumed that it is the most comprehensive (higher sampling), revised (fewer misscorings) and complete (lower amount of missing data) matrix of the genealogy. The analyses were conducted on six different genealogies under EW and IW and extended implied weighting (EIW) with a range of concavity constant values (k) between 3 and 30. Pairwise comparisons between trees were conducted using Robinson–Foulds distances normalized by the total number of groups, distortion coefficient, subtree pruning and regrafting moves, and the proportional sum of group dissimilarities. The results consistently show that IW and EIW produce results more similar to those of the last dataset than EW in the vast majority of genealogies and for all comparative measures. This is significant because almost all of these matrices were originally analysed only under EW. Implied weighting and EIW do not outperform each other unambiguously. Euclidean distances based on a principal components analysis of the comparative measures show that different ranges of k‐values retrieve the most similar results to the last generation in different genealogies. There is a significant positive linear correlation between the optimal k‐values and the number of terminals of the last generations. This could be employed to inform about the range of k‐values to be used in phylogenetic analyses based on matrix size but with the caveat that this emergent relationship still relies on a low sample size of genealogies. [ABSTRACT FROM AUTHOR]

Evidence for phylogenetic analysis comes in the form of observed similarities, and trees are selected to minimize the number of similarities that cannot be accounted for by homology (homoplasies). Thus, the classical argument for parsimony directly links homoplasy with explanatory power. When characters are hierarchically related, a first character may represent a primary structure such as tail absence/presence and a secondary (subordinate) character may describe tail colour; this makes tail colour inapplicable when tail is absent. It has been proposed that such character hierarchies should be evaluated on the same logical basis as standard characters, maximizing the number of similarities accounted for by secondary homology, i.e. common ancestry. Previous evaluations of the homology of a given ancestral reconstruction contain the unintuitive quantity "subcharacters" (number of regions of applicability). Rather than counting subcharacters, this paper proposes an equivalent but more intuitive formulation, based on counting the number of changes into each separate state. In this formulation, x‐transformations, the homoplasy for the reconstruction is simply the number of changes into the state beyond the first, summed over all states. There is thus no direct connection between homoplasy and number of steps, only between homoplasy and extra steps. The link between the two formulations is that, for any region of applicability of any character, a subcharacter can be interpreted as the change into the state that is plesiomorphic in that region. Although some authors have claimed that the equivalence between maximizing explanatory power and minimizing independent originations of similar features (i.e. the standard justification of parsimony) does not hold for inapplicable characters, evaluating homoplasy with x‐transformations clearly connects the two sides of that equation. Furthermore, as the evaluation with x‐transformations provides a direct count and a straightforward interpretation of homoplasy, it extends naturally into implied weighting, and sheds light on problems with additive, step‐matrix or continuous characters. It also allows deriving transformation costs for recoding hierarchies as step‐matrix characters (where recoded states correspond to permissible combinations of states in primary and secondary characters), so that homology of the original observations is properly measured. Those transformation costs set the cost of gaining the primary structure to the maximum difference between "present" states plus cost of loss, and difference between "present" states to the sum of user‐defined transformation costs between secondary features. With such recoding, invoking multiple independent derivations of the structure and similar features will cost as many extra "steps" as the instances of similarities (in both original characters) that are not being homologized. The step‐matrix recoding also can take into account nested dependences. We present a simple convention for naming characters, which TNT can use to automatically convert the original data into a step‐matrix form and set the proper transformation costs. Finally, the basic elements for handling inapplicable characters in the context of maximum‐likelihood inference are outlined, and some quantitative comparisons between different approaches to inapplicables are provided. [ABSTRACT FROM AUTHOR]

With approximately 3000 marine species, Tunicata represents the most disparate subtaxon of Chordata. Molecular phylogenetic studies support Tunicata as sister taxon to Craniota, rendering it pivotal to understanding craniate evolution. Although successively more molecular data have become available to resolve internal tunicate phylogenetic relationships, phenotypic data have not been utilized consistently. Herein these shortcomings are addressed by cladistically analyzing 117 phenotypic characters for 49 tunicate species comprising all higher tunicate taxa, and five craniate and cephalochordate outgroup species. In addition, a combined analysis of the phenotypic characters with 18S rDNA‐sequence data is performed in 32 OTUs. The analysis of the combined data is congruent with published molecular analyses. Successively up‐weighting phenotypic characters indicates that phenotypic data contribute disproportionally more to the resulting phylogenetic hypothesis. The strict consensus tree from the analysis of the phenotypic characters as well as the single most parsimonious tree found in the analysis of the combined dataset recover monophyletic Appendicularia as sister taxon to the remaining tunicate taxa. Thus, both datasets support the hypothesis that the last common ancestor of Tunicata was free‐living and that ascidian sessility is a derived trait within Tunicata. "Thaliacea" is found to be paraphyletic with Pyrosomatida as sister taxon to monophyletic Ascidiacea and the relationship between Doliolida and Salpida is unresolved in the analysis of morphological characters; however, the analysis of the combined data reconstructs Thaliacea as monophyletic nested within paraphyletic "Ascidiacea". Therefore, both datasets differ in the interpretation of the evolution of the complex holoplanktonic life history of thaliacean taxa. According to the phenotypic data, this evolution occurred in the plankton, whereas from the combined dataset a secondary transition into the plankton from a sessile ascidian is inferred. Besides these major differences, both analyses are in accord on many phylogenetic groupings, although both phylogenetic reconstructions invoke a high degree of homoplasy. In conclusion, this study represents the first serious attempt to utilize the potential phylogenetic information present in phenotypic characters to elucidate the inter‐relationships of this diverse marine taxon in a consistent cladistic framework. [ABSTRACT FROM AUTHOR]

Classifications in the world's tropics often involve an early and sustained adoption of Holarctic‐based patterns. Such is the case of the megadiverse subtribe Philonthina and its Neotropical (NT) members, for which generic limits are ill‐defined due to an alleged high level of homoplasy. Although a recent total‐evidence study confirmed the monophyly of a NT lineage, most of its species are assigned to the speciose genera Belonuchus Nordmann and Paederomimus Sharp, neither of them monophyletic. Here, we aim to reveal internal relationships within the NT lineage by the reassessment of characters from traditional morphology‐based systematics. Specific objectives are to test the monophyly of Belonuchus with regards to its only junior synonym, Musicoderus Sharp, as well as the placement of the six South American species of Hesperus Fauvel, a genus of Holarctic origin. We performed a phylogenetic analysis of the subtribe with focus on its NT lineage based on 132 morphological characters (50 of them novel) including 79 taxa from genera and/or species groups relevant to our study. Most novel characters assessed herein supported clades across Philonthina and its NT lineage. We found that the NT lineage diversified into at least seven clades, each of which provides a framework for future taxonomic studies. Among them, three clades containing the type species of Belonuchus, Paederomimus and Musicoderus (respectively) appear to be well supported and not closely related. The currently known South American species of Hesperus, however, are recovered within the NT lineage. We propose to resurrect Musicoderus from synonymy with Belonuchus and to transfer Hesperus novoteutonicus to Paederomimus as a new combination: Paederomimus novoteutonicus (Wendeler), comb.n. [ABSTRACT FROM AUTHOR]

Ammonite phylogeny has mainly been established based on a stratigraphic approach, with cladistics underconsidered. The main arguments against the use of cladistics are the supposed large amount of homoplasy and the small number of characters. Resolving the phylogeny of the Hildoceratidae (Early Jurassic) is especially challenging because of its large diversity and disparity. Many forms that have not been determined as closely related in previous studies exhibit very similar shapes. Moreover, some groups are morphologically very different, adding difficulties to building a unique coding scheme at a low taxonomic resolution (i.e. species). Here we propose an integrated coding scheme of the peristome shape and the ornamentation, allowing an increased level of comparison. The shape of the peristome is used as a new reference to locate ornamental features and propose new homology hypotheses. In total, 105 taxa have been analysed for 47 characters. We code continuous characters by their means and ranges ± one standard deviation. We test two weighting schemes: equal weights standardized by unit range and implied weighting with several concavity constants. This work has led to redefinition of the phylogenetic inclusivenesses of all the hildoceratid subfamilies. The new coding scheme based on peristome shapes provides the fewest homoplastic characters. The schemes appear promising to improve phylogenetic analyses in ammonoids as well as molluscs as a whole by creating a general coding framework. [ABSTRACT FROM AUTHOR]

Although 11 studies have addressed the systematics of the four families and 281 fish species of the ecomorphologically diverse Anostomoidea, none has proposed a global hypothesis of relationships. We synthesized these studies to yield a supermatrix with 463 morphological characters among 174 ingroup species, and inferred phylogeny with parsimony and Bayesian optimization. We evaluated the applicability of the supermatrix approach to morphological datasets, tested its sensitivity to missing data, determined the impact of homoplastic characters on phylogenetic resolution, and determined the distribution of homologies and homoplasies on the topology. Despite more than 60% missing data, analyses supported the monophyly of all families, and phylogenetic structure degraded only with inclusion of species with high percentages of missing data and in analyses limited to homoplasies. The latter differs modestly from the full matrix indicating phylogenetic signal in homoplastic characters. Character distributions differ across the phylogeny, with a greater prevalence of homologies at deeper nodes and homoplasies nearer the tips than expected by chance. This may suggest early diversification into distinct bauplans with subsequent diversification of faster evolving character systems. The morphological supermatrix approach is powerful and allows integration of classical data with modern methods to examine the evolution of multiple character systems. [ABSTRACT FROM AUTHOR]

Wheeler (2012) stated that minimization of ad hoc hypotheses as emphasized by Farris (1983) always leads to a preference for trivial optimizations when analysing unaligned sequence data, leaving no basis for tree choice. That is not correct. Farris's framework can be expressed as maximization of homology, a formulation that has been used to overcome the problems with inapplicables (it leads to the notion of subcharacters as a quantity to be co-minimized in parsimony analysis) and that is known not to lead to a preference for trivial optimizations when analysing unaligned sequence data. Maximization of homology, in turn, can be formulated as a minimization of ad hoc hypotheses of homoplasy in the sense of Farris, as shown here. These issues are not just theoretical but have empirical relevance. It is therefore also discussed how maximization of homology can be approximated under various weighting schemes in heuristic tree alignment programs, such as POY, that do not take into account subcharacters. Empirical analyses that use the so-called 3221 cost set (gap opening cost three, transversion and transition costs two, and gap extension cost one), the cost set that is known to be an optimal approximation under equally weighted homology in POY, are briefly reviewed. From a theoretical point of view, maximization of homology provides the general framework to understand such cost sets in terms that are biologically relevant and meaningful. Whether or not embedded in a sensitivity analysis, this is not the case for minimization of a cost that is defined in operational terms only. Neither is it the case for minimization of equally weighted transformations, a known problem that is not addressed by Kluge and Grant's (2006) proposal to invoke the anti-superfluity principle as a rationale for this minimization. [ABSTRACT FROM AUTHOR]

Several extensions to implied weighting, recently implemented in TNT, allow a better treatment of data sets combining morphological and molecular data sets, as well as those comprising large numbers of missing entries (e.g. palaeontological matrices, or combined matrices with some genes sequenced for few taxa). As there have been recent suggestions that molecular matrices may be better analysed using equal weights (rather than implied weighting), a simple way to apply implied weighting to only some characters (e.g. morphology), leaving other characters with a constant weight (e.g. molecules), is proposed. The new methods also allow weighting entire partitions according to their average homoplasy, giving each of the characters in the partition the same weight (this can be used for dynamically weighting, e.g. entire genes, or first, second, and third positions collectively). Such an approach is easily implemented in schemes like successive weighting, but in the case of implied weighting poses some particular problems. The approach has the peculiar implication that the inclusion of uninformative characters influences the results (by influencing the implied weights for the partitions). Last, the concern that characters with many missing entries may receive artificially inflated weights (because they necessarily display less homoplasy) can be solved by allowing the use of different weighting functions for different characters, in such a way that the cost of additional transformations decreases more rapidly for characters with more missing entries (thus effectively assuming that the unobserved entries are likely to also display some unobserved homoplasy). The conceptual and practical aspects of all these problems, as well as details of the implementation in TNT, are discussed. [ABSTRACT FROM AUTHOR]

Homology in cladistics is reviewed. The definition of important terms is explicated in historical context. Homology is not synonymous with synapomorphy: it includes symplesiomorphy, and Hennig clearly included both plesiomorphy and synapomorphy as types of homology. Homoplasy is error, in coding, and is analogous to residual error in simple regression. If parallelism and convergence are to be distinguished, homoplasy would be evidence of the former and analogy evidence of the latter. We discuss whether there is a difference between molecular homology and morphological homology, character state homology, nested homology (additive characters), and serial homology. We conclude by proposing a global definition of homology. ©The Will Henning Society 2011. [ABSTRACT FROM AUTHOR]

The amount of missing data in many contemporary phylogenetic analyses has substantially increased relative to previous norms, particularly in supermatrix studies that compile characters from multiple previous analyses. In such cases the missing data are non-randomly distributed and usually present in all partitions (i.e. groups of characters) sampled. Parametric methods often provide greater resolution and support than parsimony in such cases, yet this may be caused by extrapolation of branch lengths from one partition to another. In this study I use contrived and simulated examples to demonstrate that likelihood, even when applied to simple matrices with little or no homoplasy, homogeneous evolution across groups of characters, perfect model fit, and hundreds or thousands of variable characters, can provide strong support for incorrect topologies when the matrices have non-random distributions of missing data distributed across all partitions. I do so using a systematic exploration of alternative seven-taxon tree topologies and distributions of missing data in two partitions to demonstrate that these likelihood-based artefacts may occur frequently and are not shared by parsimony. I also demonstrate that Bayesian Markov chain Monte Carlo analysis is more robust to these artefacts than is likelihood. © The Willi Hennig Society 2011. [ABSTRACT FROM AUTHOR]

Here we explore the effect of missing data in phylogenetic analyses using a large number of real morphological matrices. Different percentages and patterns of missing entries were added to each matrix, and their influence was evaluated by comparing the accuracy and error of most parsimonious trees. The relationships between accuracy and error and different parameters (e.g. the number of taxa and characters, homoplasy, support) were also evaluated. Our findings, based on real matrices, agree with the simulation studies, i.e. the negative effect increases with the percentage of missing entries, and decreases with the addition of more characters. This indicates that the main problem is the lack of information, not just the presence of missing data per se. Accuracy varies with different distribution patterns of missing entries; the worst case is when missing data are concentrated in a few taxa, while the best is when the missing entries are restricted to just a few characters. The results expand our knowledge of the missing data problem, corroborate many of the findings previously published using simulations, and could be useful for empirical or theoretical studies. © The Willi Hennig Society 2009. [ABSTRACT FROM AUTHOR]

To identify a biological signal in the distribution of homoplasy, it is first necessary to isolate non-biological factors affecting its measurement. The number of states per character in a phylogenetic data matrix may indicate evolutionary flexibility and, consequently, the likelihood of recurrent evolution. However, we show here that the number of states per character limits the maximum number of steps that may be inferred using parsimony. A formula is provided for the maximum number of steps that may be taken by a character with a given number of states and taxa. We show that as more character states are included the maximum proportion of steps that can be attributed to homoplasy falls, and the greatest amount of homoplasy measurable with the consistency index declines. © The Willi Hennig Society 2009. [ABSTRACT FROM AUTHOR]

We present a molecular phylogeny of 26 out of the 28 currently accepted genera in the subfamily Chrysophylloideae (Sapotaceae) using parsimony, parsimony jackknifing, and Bayesian inference. A data matrix of 8984 characters was obtained from DNA sequences of seven chloroplast loci, two nuclear loci, indels coded as binary characters, and morphology. Our phylogenetic reconstruction suggests that Chrysophyllum, Pouteria, and Pradosia, as well as some sections within Chrysophyllum and Pouteria, are all polyphyletic. These taxa were previously described largely on the basis of unique combinations of states for a set of morphological characters. Mapping some of these characters onto one of the most parsimonious trees indicates that the symplesiomorphic flower in the subfamily was probably 5-merous, had stamens inserted in the tube orifice, staminodes, seeds with foliaceous cotyledons, exserted radicle, and endosperm. These characters have subsequently been lost multiple times and cannot be used as synapomorphies to support broad generic concepts. Despite the high degree of homoplasy some well-defined clades can be described on the basis of alternative character state combinations. Also, many of these well-supported clades appear to be restricted to particular geographical areas (e.g. all taxa in Australasia form a monophyletic group). Hence, we suggest that the segregate genera Aningeria, Malacantha, and Martiusella may ultimately be resurrected, and probably also Donella and Gambeya, but their circumscriptions are still unclear. One species, Chrysophyllum cuneifolium, may have originated from a hybridization event between continents where the maternal genome (cpDNA) comes from South America and the nuclear genome comes from Africa. © The Willi Hennig Society 2008. [ABSTRACT FROM AUTHOR]

The problem of character weighting in cladistic analysis is revisited. The finding that, in large molecular data sets, removal of third positions (with more homoplasy) decreases the number of well supported groups has been interpreted by some authors as indicating that weighting methods are unjustified. Two arguments against that interpretation are advanced. Characters that collectively determine few well-supported groups may be highly reliable when taken individually (as shown by specific examples), so that inferring greater reliability for sets of characters that lead to an increase in jackknife frequencies may not always be warranted. But even if changes in jackknife frequencies can be used to infer reliability, we demonstrate that jackknife frequencies in large molecular data sets are actually improved when downweighting characters according to their homoplasy but using properly rescaled functions (instead of the very strong standard functions, or the extreme of inclusion/exclusion); this further weakens the argument that downweighting homoplastic characters is undesirable. Last, we show that downweighting characters according to their homoplasy (using standard homoplasy-weighting methods) on 70 morphological data sets (with 50–170 taxa), produces clear increases in jackknife frequencies. The results obtained under homoplasy weighting also appear more stable than results under equal weights: adding either taxa or characters, when weighting against homoplasy, produced results more similar to original analyses (i.e., with larger numbers of groups that continue being supported after addition of taxa or characters), with similar or lower error rates (i.e., proportion of groups recovered that subsequently turn out to be incorrect). Therefore, the same argument that had been advanced against homoplasy weighting in the case of large molecular data sets is an argument in favor of such weighting in the case of morphological data sets. © The Willi Hennig Society 2008. [ABSTRACT FROM AUTHOR]

Parsimony can be related to explanatory power, either by noting that each additional requirement for a separate origin of a feature reduces the number of observed similarities that can be explained as inheritance from a common ancestor; or else by applying Popper’s formula for explanatory power together with the fact that parsimony yields maximum likelihood trees under No Common Mechanism (NCM). Despite deceptive claims made by some likelihoodists, most maximum likelihood methods cannot be justified in this way because they rely on unrealistic background assumptions. These facts have been disputed on the various grounds that ad hoc hypotheses of homoplasy are explanatory, that they are not explanatory, that character states are ontological individuals, that character data do not comprise evidence, that unrealistic theories can be used as background knowledge, that NCM is unrealistic, and that likelihoods cannot be used to evaluate explanatory power. None of these objections is even remotely well founded, and indeed most of them do not even seem to have been meant seriously, having instead been put forward merely to obstruct the development of phylogenetic methods. © The Willi Hennig Society 2008. [ABSTRACT FROM AUTHOR]

The definitions of character similarity, homology, homoplasy, heterology, parallelism and convergence are clarified in the framework of current phylogenetic methodology. They are all associated with definite patterns of character change and can consequently be tested by phylogeny building. Their crucial significance in comparative biology is illustrated using demonstrative examples. © The Willi Hennig Society 2006. [ABSTRACT FROM AUTHOR]

Traditional justifications of parsimony in phylogenetic inference assume a correspondence between character-state similarity and steps (character transformation events). In addition to similarity, justifying arguments appeal to conviction, descriptive efficiency, ad hoc hypotheses of homoplasy and frequentist probability. Each of these rationales fails in so far as the arguments are incoherent or logically inconsistent with the ontological status of what is assumed and being explained historically. An ideographic justification of parsimony, where character-states constitute transformation events, does, however, allow for a rational preference of most parsimonious phylogenetic hypotheses by invoking the anti-superfluity principle (ASP). According to ASP, explanatory power is maximized by minimizing the number of transformation events required to explain the character-states of the terminal taxa as hypotheses of homology, where the concept homology is restricted to just those inherited“ things” shared by species. The empirical significance of this rationale is exemplified with the method of direct optimization, which can identify more parsimonious hypotheses than can be confirmed with an analysis of character-state similarities. © The Willi Hennig Society 2006. [ABSTRACT FROM AUTHOR]

New examples are presented, showing that supertree methods such as matrix representation with parsimony, minimum flip trees, and compatibility analysis of the matrix representing the input trees, produce supertrees that cannot be interpreted as displaying the groups present in the majority of the input trees. These methods may produce a supertree displaying some groups present in the minority of the trees, and contradicted by the majority. Of the three methods, compatibility analysis is the least used, but it seems to be the one that differs the least from majority rule consensus. The three methods are similar in that they choose the supertree(s) that best fit the set of input trees (quantified as some measure of the fit to the matrix representation of the input trees); in the case of complete trees, it is argued that, for a supertree method to be equivalent to majority rule or frequency difference consensus, two necessary (but not sufficient) conditions must be met. First, the measure of fit between a supertree and an input tree must be symmetrical. Second, the fit for a character representing a group must be measured as absolute: either it fits or it does not fit. In the restricted case of complete and equally resolved input trees, compatibility analysis (unlike MRP and minimum flipping) fulfils these two conditions: it is symmetrical (i.e., as long as the trees have the same taxon sets and are equally resolved, the number of characters in the matrix representation of tree A that require homoplasy in tree B is always the same as the number of characters in the matrix representation of tree B that require homoplasy in tree A) and it measures fit as all-or-none. In the case of just two complete and equally resolved input trees, the two conditions (symmetry and absolute fit) are necessary and sufficient, which explains why the compatibility analysis of such trees behaves as majority consensus. With more than two such trees, these conditions are still necessary but no longer sufficient for the equivalence; in such cases, the compatibility supertree may differ significantly from the majority rule consensus, even when these conditions apply (as shown by example). MRP and minimum flipping are asymmetric and measure various degrees of fit for each character, which explains why they often behave very differently from majority rule procedures, and why they are very likely to have groups contradicted by each of the input trees, or groups supported by a minority of the input trees. © The Willi Hennig Society 2005. [ABSTRACT FROM AUTHOR]

In comparative biology, character observations initially separate similar and dissimilar characters. Only similar characters are considered for phylogeny reconstruction; their homology is attested in a two-step process, firstlya prioriof phylogeny reconstruction by accurate similarity statements, and secondlya posterioriof phylogeny analysis by congruence with other characters. Any pattern of non-homology is then a homoplasy, commonly, but vaguely, associated with“convergence”. In this logical scheme, there is no way to analyze characters which look similar, but cannot meet usual criteria for homology statements, i.e., false similarity detecteda prioriof phylogenetic analysis, even though such characters may represent evolutionarily significant patterns of character transformations. Because phylogenies are not only patterns of taxa relationships but also references for evolutionary studies, we propose to redefine the traditional concepts of parallelism and convergence to associate patterns of non-homology with explicit theoretical contexts: homoplasy is restricted to non-similarity detecteda posterioriof phylogeny analysis and related to parallelism; non-similarity detecteda prioriof phylogenetic analysis and necessarily described by different characters would then correspond to a convergence event s. str. We propose to characterize these characters as heterologous (heterology). Heterology and homoplasy correspond to different non-similarity patterns and processes; they are also associated with different patterns of taxa relationships: homoplasy can occur only in non-sister group taxa; no such limit exists for heterology. The usefulness of these terms and concepts is illustrated with patterns of acoustic evolution in ensiferan insects.© The Willi Hennig Society 2005. [ABSTRACT FROM AUTHOR]

Incongruence among trees reconstructed with different data may stem from historical (gene tree-species tree conflict) or process (character change biases) phenomena. Regardless of the source, incongruent data, as determined with “global” measures of homoplasy, have often been excluded from parsimony analysis of the combined data. Recent studies suggest that these homoplasy measures do not predict the contribution of each character to overall tree structure. Branch support measures identify, on a character to node basis, sources of support and conflict resulting from a simultaneous analysis of the data. We implement these branch support measures to identify sources of character conflict in a clade of water striders consisting of Gerris Fabricius, Aquarius Schellenberg, and Limnoporus Sta˚l species. Separate analyses of morphology, mitochondrial cytochrome oxidase I (COI), large mitochondrial ribosomal subunit (16SrRNA), and elongation factor-1α (EF-1α) data resulted in cladograms that varied in resolution and topological concordance. Simultaneous analysis of the data resulted in two trees that were unresolved for one node in a strict consensus. The topology agreed with current classification except for the placements of Aquarius chilensis and the Aquarius remigis species group closer to Gerris than to congeneric species. Branch support measures indicated that support derived from each data set varied among nodes, but COI had an overall negative effect on branch support. However, Spearman rank correlation of partitioned branch support values indicated no negative associations of branch support between any data sets and a positive association between EF-1α and 16SrRNA. Thus incongruence among data sets was not drastic and the gene-tree versus species tree phenomenon was not implicated. Biases in character change were a more likely reason for incongruence, although saturation curves and incongruence length difference for COI indicated little potential for homoplasy. However, a posteriori inspection of COI nucleotide change with reference to the simultaneous analysis tree revealed AT and codon biases. These biases were not associated with branch support measures. Therefore, it is difficult to predict incongruence or identify its cause. Exclusion of data is ill advised because every character is potentially parsimony informative. [Copyright &y& Elsevier]

Concerted homoplasy is a concern in phylogenetics because of its potential to generate strong but misleading evidence of relationships. Marques and Gnaspini (Cladistics 17, 371–381, 2001) proposed a method of recoding characters suspected of concerted homoplasy that avoids problems associated with their wholesale exclusion or inclusion. Here we show that the proposed method is analytically equivalent to excluding the recoded characters. [Copyright &y& Elsevier]

A letter to the editor is presented responding to the article "Weighting against homoplasy improves phylogenetic analysis of morphological data sets," by P.A. Goloboff, J.M. Carpenter, J.S. Arias, and D.R.M. Esquivel that appeared in issue 24, 2008 of the journal.