Preliminary comments on a phylogenetic study of the order Diphyllidea van Beneden in Carus, 1863

The basis for a preliminary analysis of the relationships within the monophyletic Diphyllidea is outlined. Information on morphological characters and their interpretation within a phylogenetic context are presented. A cladistic analysis at the species level was conducted based on a matrix of 21 morphological characters. Character polarity was determined by taxonomic outgroup analysis relative to the basal orders, Pseudophyllidea and Haplobothriidea. The phylogeny for the diphyllideans was found to be poorly resolved based on characters currently available for evaluation. Computer assisted cladistic analysis found three equally parsimonious trees with a consistency index of 0.54. The topology of these trees shows that Ditrachybothridium macrocephalum is the basal taxon and the putative sister group for species of Echinobothrium; Macrobothridium rhynchobati is grouped among species of Echinobothrium. If the classification is to be consistent with this tree, M. rhynchobati should be included in the genus Echinobothrium. This observation should be carefully examined, considering the relative paucity of useful morphological characters currently available for this group.     

How to identify (as opposed to define) a homoplasy: examples from fossil and living great apes

There is much debate on the definitions of homoplasy and homology, and on how to spot them among character states used in a phylogenetic analysis. Many advocate what I call a "processual approach," in which information on genetics, development, function, or other criteria help a priori in identifying two character states as homologous or homoplastic. I argue that the processes represented by these criteria are insufficiently known for most organisms and most characters to be reliably used to identify homoplasies and homologies. Instead, while not foolproof, phylogeny should be the ultimate test for homology. Character states are assumed to be homologous a priori because this is falsifiable and because their initial inclusion in the character-state analysis is based on the assumption that they may be phylogenetically informative. If they fall out as symplesiomorphies or synapomorphies in a phylogenetic analysis, their status as homologies remains unfalsified. If they fall out as homoplasies, having evolved independently in more than one clade, their status as homologous is falsified, and a homoplasy is identified. The character-state transformation series, functional morphology, finer levels of morphological comparison, and the distribution and correlation of characters all help to explain the presence of homoplasies in a given phylogeny. Explaining these homoplasies, and not ignoring them as "noise," should be as much a goal of phylogenetic analysis as the production of a phylogeny. Examples from the fossil record of Miocene hominoids are given to illustrate the advantages of a process-informs-pattern-recognition-after-the-fact approach to understanding the evolution of character states.     

From basepairs to birdsongs: phylogenetic data in the age of genomics

Given the quantity of molecular data now available, including complete genomes for some organisms, one can ask whether there is a need for any data beyond complete genomic sequences for phylogenetic analysis. One reason to look beyond the genome is that not all character information is encoded in organismal genomes. We propose a hierarchy of characters that ranges from biologically transmitted but nongenomically encoded characters, such as bird songs, to characters that are genomically encoded. All of these characters can retain historical information and are potentially useful for phylogenetic analysis. In addition, a number of phenotypic levels that are expressions of the genome can be identified. The question whether it is worth including any of these levels if all of the underlying sequence data have been collected arises, since issues of redundancy occur. Utilization of phenotypic levels that are ultimately based on sequences may facilitate reconstructing homologies that are not evident from sequence data alone. We propose the use of simultaneous analysis of sequence data and as many levels of phenotypic characters as possible to take advantage of homology information that may be more easily recovered from the latter. A method that eliminates redundancy to the degree that it can be detected is proposed.     

Hennig und der ursprung der tetrapoda

Hennig’s principle of analysis of characters is the best available method at present to analyse the relationship within one group or between groups. The analysis of characters has to be separated clearly from their phylogenetic and classificatory interpretation. New terms have been proposed to distinguish the status of classification of groups (synapogen, symplesiogen, paragen) from the character analysis (synapomorph etc.) and from its phylogenetic development (monophyletic, paraphyletic, polyphyletic). The origin of tetrapods is used as an example to show that every scheme of relationship depends on the use and evaluation of characters, the accepted homologies. A stabile classification is thus an illusion. No absolute criterion exists to recognize homologies.     

MORPHOMETRICS AND CLADISTICS: MEASURING PHYLOGENY IN THE SEA URCHIN ECHINOCARDIUM

A phylogenetic approach to the study of evolutionary patterns is based on taxic homologies (synapomorphies). In contrast, the recognition of evolutionary processes (namely heterochronies) involves analysis of the entire morphology. Recent developments in geometric morphometry permit analysis of morphological similarities grounded in operational homologies. Such morphometric techniques are explored (1) at the level of evolutionary processes, and (2) as a complement in exploration of phylogenetic relationships. To examplify this, we perform a two-part study of the ontogeny and phylogeny of the spatangoid sea urchin Echinocardium. First, a phylogenetic analysis of ten Recent species in the genus is performed on 18 informative characters of the test. Second, morphological divergences among the species are analyzed using procrustean (superimposition) methods based on 49 homologous points. An additive distance tree is built from a matrix of morphometric distances among adult specimens. This tree is fully congruent with the phyletic results. Ontogenetic processes are explored by inserting ontogenetic series into the analysis. A distance tree including the juvenile stages shows that the general evolutionary trend of the genus is peramorphic, but species-to-species comparisons attest that no general clinal trend exists. Our analysis emphasizes the importance of morphometric approaches in evolutionary studies (1) for the understanding of heterochronies; (2) to trace the morphological implications of phylogenetic patterns; and (3) to estimate the impact of homoplasies.     

Does minimizing homoplasy really maximize homology? MaHo: A method for evaluating homology among most parsimonious trees

Parsimony analysis aims at finding the tree that best fits hypotheses of homology. However, parsimony does not directly maximize homology, but minimizes homoplasy. When a parsimony analysis results in more than a single most-parsimonious tree (MPT), it is shown that the number of homologous characters may vary significantly. We propose a method called MaHo to identify, among the MPTs, the tree(s) that has (have) the highest number of characters that are homologies. We apply this approach to the phylogenetic relationships of the Dombeyoideae (Malvaceae) of the Mascarene Islands. A parsimony analysis was performed, including 31 representatives of the Dombeyoideae. The search resulted in 29,336 MPTs. MaHo was used in order to improve the resolution of the consensus and to increase the number of unambiguous homologies. The consensus of the 7592 MPTs presenting the highest number of homologies was chosen. This resulted in unravelling five additional synapomorphies and in reducing the number of MPTs.     

Canegrubs and cladistics: what story do adult, larval and ecological characters tell?

Abstract  Melolonthine scarabs, canegrubs, are the most important pests of Australian sugarcane. The 19 species are sometimes viewed as one pest; however, these species are quite diverse, are placed in four genera and a number of species-groupings in one of those genera, show different life histories and adult behaviours, and some have distinct soil-type preferences. Three questions concerning relationships between the canegrubs were addressed here. First, the relationships suggested by a taxonomic revision of the whole tribe based on adult characters were generally confirmed through phylogenetic analysis. Second, neither the larval characters nor the ecological characters show similar distinct phylogenetic patterns; more characters are needed to resolve the relationships. Third, the analysis of a combination of adult, larval and ecological characters, the 'total evidence' approach, does give a more robust phylogenetic arrangement than that derived from only adult characters. This analysis strengthens the understanding of the relationships among canegrubs and provides the basis for improved management systems for these pests.     

Nachtrag zu ‘Hepaticae Bomeenses (Oxford University Expedition to Sarawak, 1932)’

Abstract

Recent empirical advances in bryophyte systematics have been made in the discovery and detailed study of new characters. However, theoretical considerations have not kept pace. Proper attention has not been paid to how these new characters can be used to discover relationships among groups and produce classifications. Recent cladistic studies of the relationships of the major groups of bryophytes have demonstrated the feasibility and usefulness of an approach to phylogenetic systematics derived from the works of Hennig. A Hennigian approach attempts to find the particular hierarchical level at which a given character is a homology (i.e. a synapomorphy or shared, derived character), relationships are established by searching for congruent patterns of homologies, and the resulting hierarchical scheme is used as the basis for a classification. Such an approach provides a rigorous and explicit logic for investigating phylogeny and recognizing natural, historically discrete taxa. The theoretical advantages of a Hennigian approach are discussed and illustrated by examining the congruence of new characters data with preliminary cladograms of the major bryophyte groups.     

Integration, phylogeny, and the hominid cranial base

Basicranial features were examined in catarrhine primates and early hominids in order to demonstrate how information about morphological integration can be incorporated into phylogenetic analysis. Hypotheses purporting to explain the functional and structural relationships of basicranial characters were tested using factor analysis. Characters found to be functionally or structurally related to each other were then further examined in order to determine whether there was evidence that they were phylogenetically independent. If phylogenetic independence could not be demonstrated, then the characters were presumed to be integrated and were grouped into a complex. That complex was then treated as if it were a single character for the purposes of cladistic analysis. Factor analysis revealed that five basicranial features may be structurally related to relative brain size in hominoids. Depending on how one defines phylogenetic independence, as few as two, or as many as all of those characters might be morphologically integrated. A cladistic analysis of early hominids based on basicranial features revealed that the use of integrated complexes had a substantial effect on the phylogenetic position of Australopithecus africanus, a species whose relationships are poorly resolved. Moreover, the use of complexes also had an effect on reanalyses of certain published cladistic data sets, implying that those studies might have been biased by patterns of basicranial integration. These results demonstrate that patterns of morphological integration need to be considered carefully in all morphology-based cladistic analyses, regardless of taxon or anatomical focus. However, an important caveat is that the functional and structural hypotheses tested here predicted much higher degrees of integration than were observed. This result warns strongly that hypotheses of integration must be tested before they can be adequately employed in phylogenetic analysis. The uncritical acceptance of an untested hypothesis of integration is likely to be as disruptive to a cladistic analysis as when integration is ignored.     

Phylogenetic interpretation of ontogenetic change: sorting out the actual and artefactual in an empirical case study of centrarchid fishes

Hypothesized relationships between ontogenetic and phylogenetic change in morphological characters were empirically tested in centrarchid fishes by comparing observed patterns of character development with patterns of character evolution as inferred from a representative phylogenetic hypothesis. This phylogeny was based on 56–61 morphological characters that were polarized by outgroup comparison. Through these comparisons, evolutionary changes in character ontogeny were categorized in one of eight classes (terminal addition, terminal deletion, terminal substitution, non-terminal addition, non-terminal deletion, non-terminal substitution, ontogenetic reversal and substitution). The relative frequencies of each of these classes provided an empirical basis from which assumptions underlying hypothesized relationships between ontogeny and phylogeny were tested. In order to test hypothesized relationships between ontogeny and phylogeny that involve assumptions about the relative frequencies of terminal change (e.g. the use of ontogeny as a homology criterion), two additional phylogenies were generated in which terminal addition and terminal deletion were maximized and minimized for all characters. Character state change interpreted from these phylogenies thus represents the maxima and minima of the frequency range of terminal addition and terminal deletion for the 8.7 × 10³⁶ trees possible for centrarchids. It was found for these data that terminal change accounts for c. 75% of the character state change. This suggests either that early ontogeny is conserved in evolution or that interpretation and classification of evolutionary changes in ontogeny is biased in part by the way that characters are recognized, delimited and coded. It was found that ontogenetic interpretation is influenced by two levels of homology decision: an initial decision involving delimitation of the character (the ontogenetic sequence), and the subsequent recognition of homologous components of developmental sequences. Recognition of phylogenetic homology among individual components of developmental sequences is necessary for interpretation of evolutionary changes in ontogeny as either terminal or non-terminal. If development is the primary criterion applied in recognizing individual homologies among parts of ontogenetic sequences, the only possible interpretation of phylogenetic differences is that of terminal change. If homologies of the components cannot be ascertained, recognition of the homology of the developmental sequence as a whole will result in the interpretation of evolutionary differences as substitutions. Particularly when the objective of a study is to discover how ontogeny has evolved, criteria in addition to ontogeny must be used to recognize homology. Interpretation is also dependent upon delimitation within an ontogenetic sequence. This is in part a function of the way that an investigator ‘sees’ and codes characters. Binary and multistate characters influence interpretation differently and predictably. The use of ontogeny for determining phylogenetic polarity as previously proposed rests on the assumptions that ancestral ontogenies are conserved and that character evolution occurs predominantly through terminal addition. It was found for these data that terminal addition may comprise a maximum of 51.9% of the total character state change. It is concluded that the ontogenetic criterion is not a reliable indicator of phylogenetic polarity. Process and pattern data are collected simultaneously by those engaged in comparative morphological studies of development. The set of alternative explanatory processes is limited in the process of observing development. These form necessary starting points for the research of developmental biologists. Separating ‘empirical’ results from interpretational influences requires awareness of potential biases in the course of character selection, coding and interpretation. Consideration of the interpretational problems involved in identifying and classifying phylogenetic changes in ontogeny leads to a re-evaluation of the purpose, usefulness and information conveyed by the current classification system. It is recommended that alternative classification schemes be pursued.     

介形类系统分类与起源演化的形态、分子和化石证据

介形类(Ostracoda)因其丰富的化石记录和广布的海陆现生代表类群,而被认为是进化生物学中研究生物多样性产生机制和演变历程的颇具潜力的重要模式生物。介形类在甲壳亚门中的谱系发生位置、起源及其内部各类群间的系统关系还存在诸多争议。基于其体制构造的形态学特征,介形类被归入甲壳亚门下的颚足纲(Maxillopoda),但来自18S rDNA序列数据分析却显示Maxillopoda不是单系群。基于化石记录和壳体形态特征,高肌虫(Bradoriida)长期以来被认为是介形类的一个祖先类群,但保存有软躯体的早寒武世化石的研究表明,Bradoriida不是介形类甚至可能也不属于甲壳类。不同的研究者所强调的壳体和肢体形态特征各异,导致介形类最大的现生类群速足目(Podocopida)的四个超科之间的关系也存在诸多推测。壳体和肢体特征在系统演化意义上的不兼容,需要分子生物学等证据的介入。分子、形态和化石证据的积累及各种信息整合是系统演化研究的必然趋势。     

CLADISTIC ANALYSIS OF METAZOAN RELATIONSHIPS: A REAPPRAISAL

Abstract— A recently published cladistic analysis of metazoan relationships based on 77 embryological and morphological characters is shown to be defective with respect to both methodological issues and the interpretation of certain characters. Consequently, many conclusions of this analysis are not supported by the data. Alternative phylogenetic hypotheses are therefore proposed, based on a re-evaluation of an adapted data set.     

Phylogenetic analysis of the Pinnotheridae (Crustacea, Brachyura) based on larval morphology, with emphasis on the Dissodactylus species complex

Comparative larval morphology was used to elucidate phylogenetic relationships within the Pinnotheridae and the Dissodactylus species complex. Within the family, seven zoeal and six megalopal characters suggested two equally parsimonious phylogenetic hypotheses for pinnotherid larvae, both with Ostracotheres tridacnae representing the sister group for the Dissodactylus complex. Results indicated that the genus Pinnotheres is a polyphyletic taxon, and that the traditional subfamilial arrangement comprises paraphyletic taxa within the subfamilies Pinnotherinae and Pinnothereliinae. Certain evidence has suggested that Fabia and Juxtafubia should be excluded from the Pinnotherinae and placed into the Pinnothereliinae. Larval and adult morphology suggested that Pinnotheres politus should be included within Tumidotheres. The phylogenetic analysis within the Dissodactylus complex involved one zoeal and 16 megalopal characters. Results suggested a single phylogenetic hypothesis based on larval morphology. Combining adult morphology with larval evidence resulted in two equally parsimonious phylogenetic hypotheses, one of which agreed with a previously suggested hypothesis based only on adult characters.     

A phylogenetic analysis of Coelopidae (Diptera) based on morphological and DNA sequence data

The phylogenetic relationships of 22 species of Coelopidae are reconstructed based on a data matrix consisting of morphological and DNA sequence characters (16S rDNA, EF-1alpha). Optimal gap and transversion costs are determined via a sensitivity analysis and both equal weighting and a transversion cost of 2 are found to perform best based on taxonomic congruence, character incongruence, and tree support. The preferred phylogenetic hypothesis is fully resolved and well-supported by jackknife, bootstrap, and Bremer support values, but it is in conflict with the cladogram based on morphological characters alone. Most notably, the Coelopidae and the genus Coelopa are not monophyletic. However, partitioned Bremer Support and an analysis of node stability under different gap and transversion costs reveal that the critical clades rendering these taxa non-monophyletic are poorly supported. Furthermore, the monophyly of Coelopidae and Coelopa is not rejected in analyses using 16S rDNA that was manually aligned. The resolution of the tree based on this reduced data sets is, however, lower than for the tree based on the full data sets. Partitioned Bremer support values reveal that 16S rDNA characters provide the largest amount of tree support, but the support values are heavily dependent on analysis conditions. Problems with direct comparison of branch support values for trees derived using fixed alignments with those obtained under optimization alignment are discussed. Biogeographic history and available behavioral and genetic data are also discussed in light of this first cladogram for Coelopidae based on a quantitative phylogenetic analysis.     

Are Polyphaga (Coleoptera) really a basal neopteran lineage – a reply to Kazantsev

In a study on the mandible and mandibular articulation of larvae of the lycid genus Platerodrilus Kazantsev proposes a phylogenetic scheme with Polyphaga as a basal group of Neoptera and Lycidae as a basal group within Polyphaga. Here we point out different problems with his interpretation. The taxon sampling was not sufficient. The characters of endopterygote larvae cannot be compared to characters of adults in a phylogenetic context. The neotenic characters of female and adult male Lycidae are not sufficiently taken into account. A phylogenetic hypothesis should be based on multiple character systems and all available data must be considered. Kazantsev based his conclusions on a single isolated character complex. His hypothesis is in deep conflict with a phylogeny based on the molecular data, which clearly show that Lycidae are nested within Elateroidea. The molecular and morphological evidence also implies that females are aberrant neotenic forms and by no means ‘primitive’. Kazantsev's hypothesis is problematic because the presented data are insufficient and the character evaluation is not based on a numerical analysis.     

Phylogenetic analysis of the Amphitritinae (Polycnaeta: Terebellidae)

The phylogenetic relationships of the Amphitritinae (Polychaeta: Terebellidae) were studied using parsimony analysis of 22 external morphological characters. To choose outgroups to polarize the characters, I carried out a preliminary analysis of the relationships of the four terebellid subfamilies and the Trichobranchidae. The single most parsimonious tree from the analysis supports monophyly of the Terebellidae by the presence of ventral glandular shields. However, this character is homoplasious within the Terebellomorpha, and further evaluation of the Terebellidae is recommended. Artacama and Thelepus were chosen as outgroups for the analysis of amphitritine genera. The generic level analysis yielded seven equally parsimonious trees, which are consistent in their topologies except for the relationships among seven genera in one large clade. In all trees, Artacama is the sister taxon to a large clade within the Amphitritinae; the Artacaminae is therefore synonymized with the Amphitritinae, which is diagnosed by the presence of double rows of uncini. Within the Amphitritinae, the status of several monotypic genera is questioned; plesiomorphic character states indicated by the analysis are discussed. The results presented are offered as working hypotheses of the relationships among amphitritine genera. The large number of homoplasies indicated by the analysis emphasizes the need to further evaluate these hypotheses using additional characters. With a robust phylogenetic hypothesis of amphitritine relationships, a re-classification of the group based on apomorphic character states can be undertaken, and questions regarding the evolution of morphological characters, reproductive modes, or biogeographical patterns can be properly addressed.     

The phylogenetic position of the East Asian freshwater crayfish Cambaroides within the Northern Hemisphere Astacoidea (Crustacea, Decapoda, Astacida) based on molecular data

The phylogenetic position of the four East Asian freshwater crayfish species of the genus Cambaroides is not settled. Morphological phylogenetic analyses suggest close affinities of Cambaroides with North American Cambaridae. This view is based mainly on characters of the male and female reproductive organs. In contrast, the only molecular phylogenetic analysis so far available leaves the phylogenetic position of Cambaroides unresolved. The question of whether Cambaroides is monophyletic or paraphyletic has so far been neither addressed morphologically nor with molecular data sets. Here we present the first comprehensive phylogenetic analysis of all four currently described species of Cambaroides in the framework of Northern Hemisphere freshwater crayfish (Astacoidea) relationships based on partial sequences of two mitochondrial genes ( CO1 and 12S rRNA). Despite some evidence in favour of a monophyletic Cambaroides more data is needed to resolve this question. Our analyses suggest a close relationship of the North American Cambaridae and the European Astacidae, leaving the Asian Cambaroides basal to them. If these results hold true the similar reproductive organs of Cambaroides and American Cambaridae must be either homoplastic or ancestral for Northern Hemisphere Astacoidea.     

Somatic musculature of Tardigrada: phylogenetic signal and metameric patterns

Although studies describing molecular‐based phylogenies within tardigrades are now frequently being published, this is not the case for studies combining molecular and morphological characters. Tardigrade phylogeny is still based, from a morphological point of view, almost exclusively on chitinous structures and little attention has been given to detecting and using novel morphological data. Consequently, we analysed the musculature of seven tardigrade species belonging to the main phyletic lines by confocal laser scanning microscopy and compared these morphological results with new molecular analyses (18S+28S rRNA genes). Finally, we analysed all the data with a total evidence approach. A consilience in the phylogenetic relationships among orders and superfamilies of tardigrades was obtained among the evolutionary trees obtained from morphological, molecular and total evidence approaches. Comparative analysis on the musculature allowed the identification of serial homologies and repeated metameric patterns along the longitudinal animal body axis. A phenomenon of mosaic evolution was detected in musculature anatomy, as dorsal musculature was found to be highly modified with respect to the other body muscle groups, probably related to the evolution of dorsal cuticular plates. An understanding of tardigrade musculature anatomy will give fundamental information to understand the evolution of segmental pattern within Panarthropoda. © 2013 The Linnean Society of London     

Neurobiological constraints and fly systematics: how different types of neural characters can contribute to a higher level dipteran phylogeny

Much uncertainty still exists regarding higher level phylogenetic relationships in the insect order Diptera, and the need for independent analyses is apparent. In this paper, I present a parsimony analysis that is based on details of the nervous system of flies. Because neural characters have received little attention in modern phylogenetic analyses and the stability of neural traits has been debated, special emphasis is given to testing the robustness of the analysis itself and to evaluating how neurobiological constraints (such as levels of neural processing) influence the phylogenetic information content. The phylogenetic study is based on 14 species in three nematoceran and nine brachyceran families. All characters used in the analysis are based on anatomical details of the neural organization of the fly visual system. For the most part they relate to uniquely identifiable neurons, which are cells or cell types that can be confidently recognized as homologues among different species and thus compared. Parsimony analysis results in a phylogenetic hypothesis that favors specific previously suggested phylogenetic relationships and suggests alternatives regarding other placements. For example, several heterodactylan families (Bombyliidae, Asilidae, and Dolichopodidae) are supported in their placement as suggested by Sinclair et al. (1993), but Tipulidae and Syrphidae are placed differently. Tipulidae are placed at a derived rather than ancestral position within the Nematocera, and Syrphidae are placed within the Schizophora. The analysis suggests that neural characters generally maintain phylogenetic information well. However, by "forcing" neural characters onto conventional phylogenetic analyses it becomes apparent that not all neural centers maintain such information equally well. For example, neurons of the second-order visual neuropil, the medulla, contain stronger phylogenetic "signal" than do characters of the deeper visual center, the lobula plate. These differences may relate to different functional constraints in the two neuropils.     

Widersprüchliche Rekonstruktionen der Phylogenese am Beispiel der Ordnung Mecoptera (Schnabelfliegen; Insecta: Holometabola)

The branching patterns of phyletic lines are reconstructed by means of synapomorphic characters. Conflicting reconstructions of phylogenetic relationships result whenever symplesiomorphies or nonhomologous similarities are interpreted as synapomorphies. Two phylogenetic trees of the Mecoptera (Insecta) are taken as an example. Former reconstructions of the relationships among the mecopteran families based on wing-venation do not reflect the true genealogy, whereas the reconstruction based on the examination of the genital apparatus probably does reflect it. Not only are the genital structures highly complex but more important is the economical improvement in function of some of these structures. This may indicate the direction of evolutionary changes. The probability of false interpretation of characters in palaeontology is supposed to be higher when compared with research in neontology.