bluuebunny/crossref_metadata_2025 huggingface dataset:
The Phylogeny of the Extant Hexapod Orders
Authors: Wheeler Ward C. , Whiting Michael , Wheeler Quentin D. , Carpenter James M.
Published: July 2005

Morphological and molecular data are marshalled to address the question of hexapod ordinal relationships. The combination of 275 morphological variables, 1000 bases of the small subunit nuclear rDNA (18S), and 350 bases of the large subunit nuclear rDNA (28S) are subjected to a variety of analysis parameters (indel and transversion costs). Representatives of each hexapod order are included with most orders represented multiply. Those parameters that minimize character incongruence (ILD of Mickevich and Farris, 1981, Syst. Zool. 30, 351–370), among the morphological and molecular data sets are chosen to generate the best supported cladogram. A well‐resolved and robust cladogram of ordinal relationships is produced with the topology (Crustacea ((Chilopoda Diplopoda) ((Collembola Protura) ((Japygina Campodeina) (Archaeognatha (Zygentoma (Ephemerida (Odonata ((((Mantodea Blattaria) Isoptera) Zoraptera) ((Plecoptera Embiidina) (((Orthoptera Phasmida) (Grylloblattaria Dermaptera)) ((((Psocoptera Phthiraptera) Thysanoptera) Hemiptera) ((Neuropteroidea Coleoptera) (((((Strepsiptera Diptera) Mecoptera) Siphonaptera) (Trichoptera Lepidoptera)) Hymenoptera)))))))))))))).
Myzostomida Are Not Annelids: Molecular and Morphological Support for a Clade of Animals with Anterior Sperm Flagella
Authors: Zrzavý Jan , Hypša Václav , Tietz David F.
Published: July 2005

The myzostomes are animals with five pairs of parapodia, living as commensals or (endo)parasites mostly on crinoid and ophiuroid echinoderms. They are generally considered aberrant annelids, possibly phyllodocidan polychaetes. A phylogenetic analysis of 18S and 28S ribosomal DNA sequence data of Myzostoma glabrum, together with 60 morphological, developmental, ultrastructural, and life‐history characters, is presented to show that myzostomes are a sister group of the Cycliophora, closely related to the rotifer‐acanthocephalan clade (=Syndermata). Myzostomes and syndermates share predominantly the highly derived spermatozoa with anteriorly directed flagella (cycliophoran sperm is insufficiently known). The myzostome‐cycliophoran‐syndermate clade, accommodated within the Platyzoa (including Platyhelminthes s. str., Gastrotricha, Gnathostomulida, Syndermata, Cycliophora, and Myzostomida), is strongly supported by most analyses, regardless of alignment parameters, character combinations and weighting, species sampling, and tree‐building methods. The new name Prosomastigozoa (“forward‐flagellar animals”) is proposed for the group including three phyla (Cycliophora, Myzostomida, and Syndermata).
Parsimony with and without Scientific Justification
Authors: Kluge Arnold G.
Published: July 2005

Brower's (2000, Cladistics 16, 143–154) pursuit of a nonevolutionary cladistics, like those of others (e.g., Scotland, 2000, Syst. Biol. 49, 480–500), fails for lack of a scientific justification. His operational explication of parsimony does not necessarily rule out the use of other criteria on which to base the identification of a hierarchical branching pattern, nor does he give a compelling reason for why just that one kind of pattern is sought. In the absence of evolutionary theory, such as the descent of species, and the modification of character states, one from another, there is no scientific reason to seek congruence among character hierarchies whose origins, functions, and fates are not necessarily the same. Brower's operational parsimony is no substitute for phylogenetic parsimony, where requirements for ad hoc hypotheses of homoplasy are justifiably minimized, assuming only “descent, with modification.” In addition to maximizing explanatory power, that most parsimonious cladogram is the least disconfirmed, most highly corroborated, hypothesis.
Can Extensive Reticulation and Concerted Evolution Result in a Cladistically Structured Molecular Data Set?
Authors: Feliner Gonzalo Nieto , Aguilar Javier Fuertes , Rosselló Josep A.
Published: July 2005

Hierarchy is the main criterion for informativeness in a data set, even if no explicit reference to evolution as a causal process is provided. Sequence data (nuclear ribosomal DNA ITS) fromArmeria(Plumbaginaceae) contains a certain amount of hierarchical structure as suggested by data decisiveness and distribution of tree lengths. However, ancillary evidence suggests that extensive gene flow and biased concerted evolution in these multicopy regions have significantly shaped the ITS data set. This argument is discussed using parsimony analysis of four data sets, constructed by combining wild sequences with those from different generations of artificial hybrids (wild + F1, F2, and backcrosses; wild + backcrosses; wild + F1; wild + F2). Compared to the F1hybrids, F2show a certain degree of homogenization in polymorphic sites. This effect reduces topological disruption caused by F1and is considered to be illustrative of how extensive gene flow and biased concerted evolution may have modeled the wild ITS data. The possibility that hierarchy has arisen as a result of—or despite a significant contribution from—those two such potentially perturbing forces raises the question of what kind of signal are we recovering from this molecular data set.
Radiation of the Spider GenusDysdera(Araneae, Dysderidae) in the Canary Islands: Cladistic Assessment Based on Multiple Data Sets
Authors: Arnedo Miquel A. , Oromí Pedro , Ribera Carles
Published: July 2005

The volcanic archipelago of the Canary Islands, 100 km off the northwestern coast of Africa, harbors 43 endemic species of the mostly circum‐Mediterranean spider genusDysdera(Araneae, Dysderidae). This amounts to approximately one‐fourth of all knownDysderaspecies in an area that represents 0.1% of the range of the genus. In order to address the origin of this extraordinary number of endemic species, the phylogenetic relationships among all the endemic taxa and a sample of 27 continental species were reconstructed. A simultaneous cladistic analysis was performed on 66 morphological characters, 471 bp of the cytochrome oxidase I and 424 bp of the 16S rRNA mitochondrial genes. The preferred most parsimonious tree supports a single origin for most of the endemic species (84%), although this tree is ambiguous regarding the total number of overseas colonizations (allowing a minimum of two and a maximum of four colonization events). Our data suggest that the Canary Islands have been the source of the colonizers of some of the remaining Macaronesian archipelagoes (certainly for the Selvagem Islands and the Cape Verdes and possibly for Madeira); the Azores have been independently colonized by dysderids from the continent. The present study provides a phylogenetic framework for an exceptional case of insular species radiation, an essential tool for unraveling the factors that have promoted this amazing diversification. Species radiations in oceanic archipelagoes are excellent models for the study of speciation processes.
Support Weighting
Authors: Farris James S.
Published: July 2005

Previous weighting methods—including compatibility weighting—have assumed that homoplasy indicates unreliability, but this assumption does not seem to hold for large molecular data matrices. Reliability can be better assessed by support weighting, which measures the degree to which the changes in a character (site) are concentrated in the supported branches of a tree. Jackknife resampling can be used to generate randomly selected suites of initial weights in successive support weighting, and this provides a way of assessing the stability of successive weighting results.
Phylogeny and Systematic Position of Opiliones: A Combined Analysis of Chelicerate Relationships Using Morphological and Molecular Data1
Authors: Giribet Gonzalo , Edgecombe Gregory D. , Wheeler Ward C. , Babbitt Courtney
Published: July 2005

The ordinal level phylogeny of the Arachnida and the suprafamilial level phylogeny of the Opiliones were studied on the basis of a combined analysis of 253 morphological characters, the complete sequence of the 18S rRNA gene, and the D3 region of the 28S rRNA gene. Molecular data were collected for 63 terminal taxa. Morphological data were collected for 35 exemplar taxa of Opiliones, but groundplans were applied to some of the remaining chelicerate groups. Six extinct terminals, including Paleozoic scorpions, are scored for morphological characters. The data were analyzed using strict parsimony for the morphological data matrix and via direct optimization for the molecular and combined data matrices. A sensitivity analysis of 15 parameter sets was undertaken, and character congruence was used as the optimality criterion to choose among competing hypotheses. The results obtained are unstable for the high‐level chelicerate relationships (except for Tetrapulmonata, Pedipalpi, and Camarostomata), and the sister group of the Opiliones is not clearly established, although the monophyly of Dromopoda is supported under many parameter sets. However, the internal phylogeny of the Opiliones is robust to parameter choice and allows the discarding of previous hypotheses of opilionid phylogeny such as the “Cyphopalpatores” or “Palpatores.” The topology obtained is congruent with the previous hypothesis of “Palpatores” paraphyly as follows: (Cyphophthalmi (Eupnoi (Dyspnoi + Laniatores))). Resolution within the Eupnoi, Dyspnoi, and Laniatores (the latter two united as Dyspnolaniatores nov.) is also stable to the superfamily level, permitting a new classification system for the Opiliones.
Testing Methods: The Evaluation of Discovery Operations in Evolutionary Biology
Authors: Grant Taran
Published: July 2005

Scientific discovery requires both abstract, theoretically defined concepts and discovery operations formed by sets of rules that permit the empirical detection of instances of those concepts. In this paper, I examine the ontological status of discovery operations and the tests employed to evaluate them in evolutionary biology. Attention is drawn to the distinction between nomothetic (universal, predictive) and ideographic (historical, retrodictive) discovery operations, and between complementary and exclusive discovery operations. Three types of tests of discovery operations are commonly employed in evolutionary biology. Theoretical tests aim to show that a discovery operation is inconsistent with accepted, well‐corroborated, empirical theories. Empirical tests evaluate the performance of competing discovery operations in terms of their results when applied to the same empirical data sets. Philosophical tests aim to show that an operation is inconsistent with logical and epistemological principles. Appropriately designed theoretical and philosophical tests of ideographic discovery operations may be scientifically valid. Empirical tests, however, are incapable of evaluating the scientific merits of competing discovery operations. Nonetheless, empirical comparisons (not tests) of competing discovery operations may provide insight into the ways discovery operations may be misleading and therefore may play an important role in stimulating critical debate and eventually establishing a scientifically optimal operation. In practice, theoretical and philosophical tests are often combined to test competing discovery operations as rigorously as possible.
Molecular Rates Parallel Diversification Contrasts between Carnivorous Plant Sister Lineages1
Authors: Jobson Richard W. , Albert Victor A.
Published: July 2005

In the carnivorous plant family Lentibulariaceae, the bladderwort lineage (Utricularia and Genlisea) is substantially more species‐rich and morphologically divergent than its sister lineage, the butterworts (Pinguicula). Bladderworts have a relaxed body plan that has permitted the evolution of terrestrial, epiphytic, and aquatic forms that capture prey in intricately designed suction bladders or corkscrew‐shaped lobster‐pot traps. In contrast, the flypaper‐trapping butterworts maintain vegetative structures typical of angiosperms. We found that bladderwort genomes evolve significantly faster across seven loci (the trnL intron, the second trnL exon, the trnL–F intergenic spacer, the rps16 intron, rbcL, coxI, and 5.8S rDNA) representing all three genomic compartments. Generation time differences did not show a significant association. We relate these findings to the contested speciation rate hypothesis, which postulates a relationship between increased nucleotide substitution and increased cladogenesis.
Behavioral Phylogeny of Corbiculate Apidae (Hymenoptera; Apinae), with Special Reference to Social Behavior
Authors: Noll Fernando B.
Published: July 2005

The phylogenetic relationships among the four tribes of corbiculate bees (Euglossini, Bombini, Meliponini, and Apini) are controversial. There is substantial incongruence between morphological and molecular data, and the single origin of eusociality is questionable. The use of behavioral characters by previous workers has been restricted to some typological definitions, such as solitary and eusocial. Here, I expand the term “social” to 42 characters and present a tree based only on behavioral characters. The reconstructed relationships were similar to those observed in morphological and “total evidence” analyses, i.e., Euglossini + (Bombini + (Meliponini + Apini)), all of which support a single origin of eusociality.
Conflict between Amino Acid and Nucleotide Characters
Authors: Simmons Mark P. , Ochoterena Helga , Freudenstein John V.
Published: July 2005

Slowly evolving characters, such as amino acids and replacement substitutions, have generally been favored over faster evolving characters for inferring phylogenetic relationships. However, amino acids constitute composite characters and, because of the degenerate genetic code, are subject to convergence. Based on an analysis of atpB and rbcL in 567 seed plants, we show that silent substitutions may be more phylogenetically informative than replacement substitutions and that artifacts caused by composite characters and/or convergence cause clades on amino acid trees to conflict with nucleotide trees and independent evidence. These findings indicate that coding nucleotide sequences only as amino acid characters for phylogenetic analysis provides little benefit and may yield misleading results.
The Palaeoptera Problem: Basal Pterygote Phylogeny Inferred from 18S and 28S rDNA Sequences
Authors: Hovmöller Rasmus , Pape Thomas , Källersjö Mari
Published: July 2005

Monophyly of the pterygote insects is generally accepted, but the relationships among the three basal branches (Odonata, Ephemeroptera and Neoptera) remain controversial. The traditional view, to separate the pterygote insects in Palaeoptera (Odonata + Ephemeroptera) and Neoptera, based on the ability or inability to fold the wings over the abdomen, has been questioned. Various authors have used different sets of morphological characters in support of all three possible arrangements of the basal pterygote branches. We sequenced 18S and 28S rDNA from 18 species of Odonata, 8 species of Ephemeroptera, 2 species of Neoptera, and 1 species of Archaeognatha in our study. The new sequences, in combination with sequences from GenBank, have been used in a parsimony jackknife analysis resulting in strong support for a monophyletic Palaeoptera. Morphological evidence and the phylogenetic implications for understanding the origin of insect flight are discussed.
On RASA
Authors: Faivovich Julián
Published: July 2005

Relative Apparent Synapomorphy Analysis (RASA) was recently proposed as a way to measure phylogenetic signal, choose “optimal” outgroups, find long branches, and eliminate long‐branch attraction. In this paper it is shown with simple examples that RASA has several problems. The null regression model used by RASA to measure phylogenetic signal does not have a straightforward relation to phylogenetic information. RASA detects long branches, but does not discriminate between long branches that mislead an analysis and those that do not. Rooted RASA, which is used for “optimal outgroup analysis,” is shown to be an inappropriate measure of “+esiomorphy content”.
Artifacts of Coding Amino Acids and Other Composite Characters for Phylogenetic Analysis
Authors: Simmons Mark P. , Freudenstein John V.
Published: July 2005

A phylogenetic analysis can be no better than the characters on which it is based. Just as it is inappropriate to code character states of individual characters as separate presence/absence characters, it is inappropriate to combine independent characters because not all information in the data is being utilized. Composite characters link otherwise discernible states from different characters together to form new character states. There are two related problems with this coding. First, there is a loss of hierarchic information between the reductive and composite characters when unordered states are used. Second, the linking of separate characters that occurs during the construction of composite character states can create putative synapomorphies that were not present in the separate characters. For amino acid characters, the problem may occur whenever more than one position of a codon is variable among the terminals sampled. Groups that are resolved as paraphyletic with reductive coding may be resolved as monophyletic with composite coding. The artificial character states indicated by the amino acid characters are unlikely to be congruent with the true gene tree.
Partitioned Bremer support and multiple trees
Authors: Lambkin Christine L. , Lee Michael S.Y. , Winterton Shaun L. , Yeates David K.
Published: July 2005

Partitioned Bremer support (PBS) is a valuable means of assessing congruence in combined data sets, but some aspects require clarification. When more than one equally parsimonious tree is found during the constrained search for trees lacking the node of interest, averaging PBS for each data set across these trees can conceal conflict, and PBS should ideally be examined for each constrained tree. Similarly, when multiple most parsimonious trees (MPTs) are generated during analysis of the combined data, PBS is usually calculated on the consensus tree. However, extra information can be obtained if PBS is calculated on each of the MPTs or even suboptimal trees.
Taxon sampling and seed plant phylogeny
Authors: Rydin Catarina , Källersjö Mari
Published: July 2005

We investigated the effects of taxon sampling on phylogenetic inference by exchanging terminals in two sizes of rbcL matrices for seed plants, applying parsimony and bayesian analyses to ten 38‐taxon matrices and ten 80‐taxon matrices. In comparing tree topologies we concentrated on the position of the Gnetales, an important group whose placement has long been disputed. With either method, trees obtained from different taxon samples could be mutually contradictory and even disagree on groups that seemed strongly supported. Adding terminals improved the consistency of results for unweighted parsimony, but not for parsimony with third positions excluded and not for bayesian analysis, particularly when the general time‐reversible model was employed. This suggests that attempting to resolve deep relationships using only a few taxa can lead to spurious conclusions, groupings unlikely to be repeatable with different taxon samplings or larger data sets. The effect of taxon sampling has not generally been recognized, and phylogenetic studies of seed plants have often been based on few taxa. Such insufficient sampling may help explain the variety of phylogenetic hypotheses for seed plants proposed in recent years. We recommend that restricted data sets such as single‐gene subsets of multigene studies should be reanalyzed with alternative selections of terminals to assess topological consistency.
Sensitivity analysis of different methods of coding taxonomic polymorphism: an example from higher‐level bat phylogeny
Authors: Simmons Nancy B. , Geisler Jonathan H.
Published: July 2005

New information concerning strengths and weaknesses of different methods of coding taxonomic polymorphisms suggests that results of some previous studies may have been unintentionally biased by the methods employed. In this study, we demonstrate that a form of sensitivity analysis can be used to evaluate the effects of different methods of coding taxonomic polymorphisms on the outcome of phylogenetic analyses. Our earlier analysis of higher‐level relationships of bats (Mammalia: Chiroptera) employed superspecific taxa as terminals and scored taxonomic polymorphisms using ambiguity coding. Application of other methods of dealing with polymorphisms (excluding variable characters, inferring ancestral states, majority coding) to the same data yields phylogenetic results that differ somewhat from those originally reported based on ambiguity coding. Monophyly of some clades was supported in all analyses (e.g., Microchiroptera, Rhinopomatoidea, and Nataloidea), while other groups found to be monophyletic in the original study (e.g., neotropical Nataloidea) appeared unresolved or nonmonophyletic when other methods were used to code taxonomic polymorphisms. Several groupings that were apparently refuted in the initial study (e.g., Noctilionoidea including Mystacinidae) were supported in some analyses, reducing some of the apparent incongruence between the trees in our earlier analysis (which were based principally on morphology) and other trees based on molecular data. Perceived support for various groupings (branch support, bootstrap values) were in some cases significantly affected by the methods employed. These results indicate that sensitivity analysis provides a useful tool for evaluating effects of different methods of dealing with taxonomic polymorphism in superspecific terminal taxa. Variation in results obtained with different methods suggests that it is always preferable to sample at the species level when higher‐level taxa exhibit taxonomic polymorphism, thus avoiding methodological biases associated with different methods of dealing with taxonomic polymorphisms during data analysis.
Parsimony, likelihood, and simplicity
Authors: Goloboff Pablo A.
Published: July 2005

The latest charge against parsimony in phylogenetic inference is that it involves estimating too many parameters. The charge is derived from the fact that, when each character is allowed a branch length vector of its own (instead of the homogeneous branch lengths assumed in current likelihood models), the results for likelihood and parsimony are identical. Parsimony, however, can also be derived from simpler models, involving fewer parameters. Therefore, parsimony provides (as many authors had argued before) the simplest explanation of the data, or the most realistic, depending on one's views. If (as argued by likelihoodists) phylogenetic inference is to use the simplest model that provides sufficient explanation of the data, the starting point of phylogenetic analyses should be parsimony, not maximum likelihood. If the addition of new parameters (which increase the likelihood) to a parsimony estimation is seen as desirable, this may lead to a preference for results based on current likelihood models. If the addition of parameters is continued, however, the results will eventually come back to the same place where they had started, since allowing each character a branch length of its own also produces parsimony. Parsimony can be justified by very different types of models—either very complex or very simple. This suggests that parsimony does have a unique place among methods of phylogenetic estimation.
Cladistic analysis of languages: Indo‐European classification based on lexicostatistical data
Authors: Rexová Kateřina , Frynta Daniel , Zrzavý Jan
Published: July 2005

The phylogeny of the Indo‐European (IE) language family is reconstructed by application of the cladistic methodology to the lexicostatistical dataset collected by Dyen (about 200 meanings, 84 speech varieties, the Hittite language used as a functional outgroup). Three different methods of character coding provide trees that show: (a) the presence of four groups, viz., Balto‐Slavonic clade, Romano‐Germano‐Celtic clade, Armenian‐Greek group, and Indo‐Iranian group (the two last groups possibly paraphyletic); (b) the unstable position of the Albanian language; (c) the unstable pattern of the basalmost IE differentiation; but (d) the probable existence of the Balto‐Slavonic–Indo‐Iranian (“satem”) and the Romano‐Germano‐Celtic (+Albanian?) superclades. The results are compared with the phenetic approach to lexicostatistical data, the results of which are significantly less informative concerning the basal pattern. The results suggest a predominantly branching pattern of the basic vocabulary phylogeny and little borrowing of individual words. Different scenarios of IE differentiation based on archaeological and genetic information are discussed.
Taxon names as paradigms: the structure of nomenclatural revolutions
Authors: Härlin Mikael
Published: July 2005

In the present paper I argue that the two systems of phylogenetic nomenclature hitherto proposed represent, in a generalized sense, two different philosophies for how science develops and progresses. The phylogenetic system of definition initially proposed by de Queiroz and Gauthier [Syst. Zool. 39 (1990) 307], and later labeled PSD, is typically Popperian in the sense that science progresses toward truth by an accumulation of knowledge. Phylogenetic definitions of taxon names are assumed to adapt automatically to each new hypothesis of phylogeny, thereby reflecting better and better hypotheses. The phylogenetic system of reference proposed by Härlin [Zool. Scr. 27 (1998a) 381], on the other hand, is more Kuhnian, because it is built on the idea that successive hypotheses are incommensurable (and thus not cumulative) and that taxon names might be equalled with low‐level paradigms.
On the relationship between content, ancestor, and ancestry in phylogenetic nomenclature
Authors: Härlin Mikael
Published: July 2005

In this paper I draw attention to the concepts of content and ancestry in phylogenetic nomenclature. I argue that these concepts are tightly linked and that they cannot be separated as suggested by Bryant and Cantino [Biol. Rev. 77 (2002) 39] in their recent response to a critique of phylogenetic nomenclature. In addition, I argue that the basic assumption in phylogenetic nomenclature that a taxon name always refers to the same ancestor or ancestry is questionable.
A morphological analysis of marsupial mammal higher‐level phylogenetic relationships
Authors: Horovitz Inés , Sánchez‐Villagra Marcelo R
Published: July 2005

Phylogenetic relationships among marsupial taxa have proven to be more complex than the simple grouping of species by continent. Recent marsupials are distributed across the New World, Australia, New Guinea, and certain neighboring islands. Morphological characteristics of various groups bridge different geographical areas. We investigated the origin of these characteristics by assembling a morphological data matrix consisting of a new suite of 149 postcranial characters and incorporated a series of previously published data on the craniodental (76 characters) and soft tissue (5 characters) anatomy. Twenty‐one marsupial terminal taxa representing all the major radiations of marsupials and 10 outgroups, most of which are exceptionally well‐preserved fossils such as Vincelestes, Ukhaatherium, and a few basal metatherian taxa, were investigated. A maximum parsimony analysis was conducted, resulting in one most parsimonious tree. Relationships among outgroups are congruent with current understanding of mammalian phylogeny. All currently accepted marsupial orders were recovered by the analysis. We confirmed previous results showing the South American “monito del monte”Dromiciops nested within the Australasian radiation. Within this australidelphian clade, Dromiciops was closely allied with the Diprotodontia. The South American paucituberculates appeared more closely related to the Australidelphia than to the American Didelphimorphia. The marsupial mole Notoryctes and the Peramelia were closely allied to each other and in turn were the sister group of the Dromiciops plus Diprotodontia clade. This pattern of relationships left Dasyuromorphia as the most basal offshoot of the Australidelphia. Whereas this tree topology recovers some signal that had been detected by previous studies, morphological and/or molecular, some novel hypotheses are also supported.
Morphology versus molecules: resolution of the positions of Nymphalis, Polygonia, and related genera (Lepidoptera: Nymphalidae)
Authors: Wahlberg Niklas , Nylin Sören
Published: July 2005

The debate on whether to combine different data sets for simultaneous analysis has continued to the present day unabated. We have studied the effects of combining one morphological data set with four molecular data sets (two mitochondrial gene sequences and two nuclear gene sequences) for a group of butterflies belonging to the tribe Nymphalini using partitioned Bremer support. We particularly focus our attention on a group of species belonging to the genera Aglais, Inachis, Roddia, Nymphalis, Kaniska, and Polygonia. We find that, despite significant incongruence between most data partitions, all data partitions contribute positively to the support of most nodes in the most parsimonious trees found for the combined data set. We also find that the morphological data set resolves one particular node (Kaniska basal to Polygonia) with good support, while the molecular data sets are ambiguous about the existence of this node. We suggest that partitioned Bremer support allows one to critically appraise the robustness of each node in a given tree and thereby identify nodes that may change with the addition of new data and nodes that are likely to remain unchanged with new data. We also suggest that morphological data are still crucial to our being able to understand the relationships of extant organisms, despite published views to the contrary. Based on our results we suggest that Inachis should be synonymized with Aglais, Roddia with Nymphalis, and Kaniska with Polygonia.
Molecular phylogeny of the Sphaerophorus globosus species complex
Authors: Högnabba Filip , Wedin Mats
Published: July 2005

The Sphaerophorus globosus complex (Lecanorales, lichenized Ascomycota) shows a large morphological variation, and three relatively distinct morphotypes can be distinguished in parts of the distribution area. Here, we utilize a multigene‐based maximum‐parsimony approach (nITS+
LSU rDNA, mtSSU rDNA, β‐tubulin, and actin) to investigate whether these morphotypes constitute distinct species. The results show that there are at least two well‐supported monophyletic groups that we interpret as phylogenetic species within the S. globosus complex. These species do not completely correspond to the predefined morphotypes. One group, an apparently undescribed species, contains noncoralloid specimens from the North American Pacific Northwest. The other group, S. globosus, consists of two well‐supported monophyletic groups: one contains coralloid epiphytic specimens from the North American Pacific Northwest that are morphologically indistinguishable from epiphytic specimens from Europe and are presently interpreted as belonging to the same species and the other is morphologically variable and contains terrestrial specimens from Europe, North America, and southernmost South America and coralloid epiphytic and epilithic specimens from Europe. The results suggest that the population in southernmost South America originated by long‐distance dispersal from arctic populations in the Northern Hemisphere.
A priori and a posteriori methods in comparative evolutionary studies of host–parasite associations
Authors: Dowling Ashley P.G , van Veller Marco G.P , Hoberg Eric P , Brooks Daniel R
Published: July 2005

Brooks parsimony analysis (BPA) and reconciliation methods in studies of host–parasite associations differ fundamentally, despite using the same null hypothesis. Reconciliation methods may eliminate or modify input data to maximize fit of single parasite clades to a null hypothesis of cospeciation, by invoking different a priori assumptions, including a known host phylogeny. By examining the degree of phylogenetic congruence among multiple parasite clades, using hosts as analogs of taxa but not presuming a host phylogeny or any degree of cospeciation a priori, BPA modifies the null hypothesis of cospeciation if necessary to maintain the integrity of the input data. Two exemplars illustrate critical empirical differences between reconciliation methods and BPA: (1) reconciliation methods rather than BPA may select the incorrect general host cladogram for a set of data from different clades of parasites, (2) BPA rather than reconciliation methods provides the most parsimonious interpretation of all available data, and (3) secondary BPA, proposed in 1990, when applied to data sets in which host‐switching produces hosts with reticulate histories, provides the most parsimonious and biologically realistic interpretations of general host cladograms. The extent to which these general host cladograms, based on cospeciation among different parasite clades inhabiting the same hosts, correspond to host phylogeny can be tested, a posteriori, by comparison with a host phylogeny generated from nonparasite data. These observations lead to the conclusion that BPA and reconciliation methods are designed to implement different research programs based on different epistemologies. BPA is an a posteriori method that is designed to assess the host context of parasite speciation events, whereas reconciliation methods are a priori methods that are designed to fit parasite phylogenies to a host phylogeny. Host‐switching events are essential for explaining complex histories of host–parasite associations. BPA assumes coevolutionary complexity (historical contingency), relying on parsimony as an a posteriori explanatory tool to summarize complex results, whereas reconciliation methods, which embody formalized assumptions of maximum cospeciation, are based on a priori conceptual parsimony. Modifications of basic reconciliation methods, embodied in TreeMap 1.0 and TreeMap 2.02, represent the addition of weighting schemes in which the researcher specifies allowed departures from cospeciation a priori, with the result that TreeMap results more closely agree with BPA results than do reconciled tree analysis results.
The evolution of the laterophysic connection with a revised phylogeny and taxonomy of butterflyfishes (Teleostei: Chaetodontidae)
Authors: Smith Wm.Leo , Webb Jacqueline F , Blum Stanley D
Published: July 2005

The higher‐level relationships of butterflyfishes were examined using 37 morphological characters. This analysis combines characters derived from a histological study describing variation in the morphology of the laterophysic connection (an association between the swim bladder and the lateral‐line canals) with previously described morphological characters. The phylogenetic analysis resulted in four equally parsimonious trees that only differed in the placement of two of the 11 chaetodontid genera (Amphichaetodon and Forcipiger). We compare our analysis with previous hypotheses, present a new taxonomy consistent with the proposed cladistic relationships, and diagnose Chaetodon with five unreversed synapomorphies, including the evolution of characters composing the laterophysic connection. A new character‐based diagnosis of Chaetodon is provided and species are allocated accordingly; Chaetodon now includes the former Parachaetodon ocellatus and excludes the former subgenera Prognathodes and Roa. The evolution of the laterophysic connection is examined by optimizing character‐state transformations on the new hypothesis of relationships.
Phylogenetics of Sigmodontinae (Rodentia, Muroidea, Cricetidae), with special reference to the akodont group, and with additional comments on historical biogeography
Authors: D’Elıacute;a Guillermo
Published: July 2005

This is the first cladistic analysis of sigmodontine rodents (Cricetidae, Sigmodontinae) based on nuclear and mitochondrial DNA sequences. Two most parsimonious cladograms (7410 steps in length; CI=0.199; RI=0.523) were discovered. Sigmodontinae appears well supported. Sigmodon is sister to the remaining living sigmodontines. It is shown that Euneomys is not a phyllotine and that the Reithrodon group is not monophyletic. Results corroborate that the abrothricines form a natural group that is not part of the akodont radiation. The akodontine tribe is well supported, and is composed of five main clades, whose limits and relationships are thoroughly discussed. For instance, the scapteromyines do not form a natural group and they fall within the akodontine clade. Additionally, I present some taxonomic judgments and comments on the historical biogeography of sigmodontines in the light of the newly discovered relationships. For example, five akodontine divisions are suggested: the Akodon, the Bibimys, the Blarinomys, the Oxymycterus, and the Scapteromys Divisions. It is shown that traditional hypotheses of sigmodontine historical biogeography are falsified by the recovered topology.
Improvements to resampling measures of group support
Authors: Goloboff Pablo A , Farris James S , Källersjö Mari , Oxelman Bengt , Ramıacute;rez Martıacute;n J , Szumik Claudia A
Published: July 2005

Several aspects of current resampling methods to assess group support are reviewed. When the characters have different prior weights or some state transformation costs are different, the frequencies under either bootstrapping or jackknifing can be distorted, producing either under‐ or overestimations of the actual group support. This is avoided by symmetric resampling, where the probability p of increasing the weight of a character equals the probability of decreasing it. Problems with interpreting absolute group frequencies as a measure of the support are discussed; group support does not necessarily vary with the frequency itself, since in some cases groups with positive support may have much lower frequencies than groups with no support at all. Three possible solutions for this problem are suggested. The first is measuring the support as the difference in frequency between the group and its most frequent contradictory group. The second is calculating frequencies for values of p below the threshold under which the frequency ranks the groups in the right order of support (this threshold may vary from data set to data set). The third is estimating the support by using the slope of the frequency as a function of different (low) values of p; when p is low, groups with actual support have negative slopes (closer to 0 when the support is higher), and groups with no support have positive slopes (larger when evidence for and against the group is more abundant).
From Haeckel to Hennig: the early development of phylogenetics in German‐speaking Europe
Authors: Willmann Rainer
Published: July 2005

An outline of the development of phylogenetic thinking and methodology in German literature published between 1862 and 1942 is presented. Central European biologists and palaeontologists of the first post‐Darwinian generation of biologists holding evolutionary views were directly stimulated by Darwin. Members of the second generation, mostly born after 1850, were largely influenced also by colleagues of the first post‐Darwinian generation, mainly by Haeckel. Among them were O. Abel, V. Franz, R. Hertwig, A. Naef, L. Plate, and R. v. Wettstein. Opinions on the relationship between systematics and phylogeny differed considerably. Many authors admitted that phylogeny must be mirrored in systematics but at the same time shared Haeckel's views on classification, which permitted paraphyletic groupings. Particularly Abel and Naef took systematics several steps further, and many important elements of phylogenetic systematics were developed several decades before Hennig. Naef presented a definition of a phylogenetic group that exactly matches Hennig's definition of monophyly. He also formulated a species concept that was implicitly based on reproductive isolation. This was an important presupposition for viewing speciation as the splitting of a stem species into daughter species. However, many authors of the first half of the 20th century repeated old, but established views on phylogenetics, while others overlooked or misunderstood earlier progressive views thus causing slow development of phylogenetic systematics in Central Europe. Its development almost stopped between 1925 and 1950, because of a widespread shift towards typology and extreme idealistic morphology. During that time very few persons such as W. Zimmermann and W. Hennig assembled elements of phylogenetic systematics and combined them with their own thoughts to create a sound theory and methodology.
Sources of character conflict in a clade of water striders (Heteroptera: Gerridae)
Authors: Damgaard Jakob , Cognato Anthony I
Published: July 2005

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 Stål 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.
The deep divergences of neornithine birds: a phylogenetic analysis of morphological characters
Authors: Mayr Gerald , Clarke Julia
Published: July 2005

Consensus is elusive regarding the phylogenetic relationships among neornithine (crown clade) birds. The ongoing debate over their deep divergences is despite recent increases in available molecular sequence data and the publication of several larger morphological data sets. In the present study, the phylogenetic relationships among 43 neornithine higher taxa are addressed using a data set of 148 osteological and soft tissue characters, which is one of the largest to date. The Mesozoic non‐neornithine birds Apsaravis, Hesperornis, and Ichthyornis are used as outgroup taxa for this analysis. Thus, for the first time, a broad array of morphological characters (including both cranial and postcranial characters) are analyzed for an ingroup densely sampling Neornithes, with crown clade outgroups used to polarize these characters. The strict consensus cladogram of two most parsimonious trees resultant from 1000 replicate heuristic searches (random stepwise addition, tree‐bisection‐reconnection) recovered several previously identified clades; the at‐one‐time contentious clades Galloanseres (waterfowl, fowl, and allies) and Palaeognathae were supported. Most notably, our analysis recovered monophyly of Neoaves, i.e., all neognathous birds to the exclusion of the Galloanseres, although this clade was weakly supported. The recently proposed sister taxon relationship between Steatornithidae (oilbird) and Trogonidae (trogons) was recovered. The traditional taxon “Falconiformes” (Cathartidae, Sagittariidae, Accipitridae, and Falconidae) was not found to be monophyletic, as Strigiformes (owls) are placed as the sister taxon of (Falconidae + Accipitridae). Monophyly of the traditional “Gruiformes” (cranes and allies) and ”Ciconiiformes” (storks and allies) was also not recovered. The primary analysis resulted in support for a sister group relationship between Gaviidae (loons) and Podicipedidae (grebes)—foot‐propelled diving birds that share many features of the pelvis and hind limb. Exclusion of Gaviidae and reanalysis of the data set, however, recovered the sister group relationship between Phoenicopteridae (flamingos) and grebes recently proposed from molecular sequence data.
Brooks Parsimony Analysis: a valiant failure
Authors: Siddall Mark E , Perkins Susan L
Published: July 2005

A recent comparison of two methods for examining correlated host and parasite phylogenies, namely TreeMap 1.0 and Brooks Parsimony Analysis concluded that the latter method performed better and is to be preferred. Reevaluation of the examples contrived for that study demonstrates that the two methods were only compared on one kind of problem (widespread parasite) for which there is an easy fix for TreeMap 1.0. Other kinds of problems like host‐switching among sister taxa or host‐switching over great distances across a host tree befuddle BPA even as they are readily resolved parsimoniously by TreeMap 1.0. These difficulties, compounded with inaccurate counting of ad hoc hypotheses required by its solutions render BPA unsuitable for comparison of host and parasite phylogenies.
The origin of photic behavior and the evolution of sexual communication in fireflies (Coleoptera: Lampyridae)
Authors: Branham Marc A , Wenzel John W
Published: July 2005

Through a phylogenetic analysis using adult morphological characters, we show that the origin of bioluminescence in cantharoid beetles appears to predate the origin of the family Lampyridae. The ability to produce and emit photic signals was first gained by larvae and appears to function as an aposematic warning display; it was subsequently gained in adults and is used as a sexual signal. Our analysis also suggests that while pheromonal sexual signals are used basally in the family, they are used in conjunction with and then subsequently replaced by photic signals in some lampyrid lineages. Both photic signals and the photic organs used to produce them have become greatly elaborated in the fireflies that no longer employ pheromonal sexual signals. In addition, the ability to produce a flashed sexual signal appears to have arisen at least three times in the family Lampyridae. Convergent evolution is also evident in a number of adult male photic organ morphologies. Further, we recommend that individual signal system components be compared rather than overall signal system complexity. The use of this strategy may allow one to recognize and better interpret adaptive correlations despite convergence or loss. We demonstrate that phylogenetic analysis is a powerful tool even for rapidly evolving traits.