Phylogeny and evolutionary biology

The phylogenetic system of Hennig, which was designed for classification of synchronous organisms, has only been adapted secondarily to total reconstruction of phylogeny. All fields of fundamental biology are related to the development of evolutionary theory. A better understanding of the origin of life requires new concepts of the historical geology of prebiological environment and new concepts of molecular genetics and biochemistry concerning ribonucleic acids as the initial units for the origin of life.     

Hennigs theorem und die strategie des stammesgeschichtlichen rekonstruierens die agnathen-gnathostomen-beziehung als beispiel

The heavily disputed methodology for the formulation of cladograms advocated by Hennig is subjected to a strict test as far as theoretical consistency and applicability are concerned. It can be convincingly shown that Hennig’s theorem contains indispensible postulates as it requires the establishment of plesiomorphic and apomorphic situations in the process of reconstructing the phylogenetic connections between existing fossil or recent organisms. Hennig’s view that fossil remains cannot by themselves disclose the phylogenetic interrelationships of the organisms and require an assessement of the characters is supported by the model for the evolution of the jaw apparatus in lower vertebrates. The model that is based on the main tenets of an approach for reconstructing phylogenetic transformations provides the key for the evolutionary position of fossil fish groups. In contrast to the logical and theoretical clarity Hennig’s approach does not offer any conclusive arguments as how to discriminate plesiomorphic and apomorphic character states and by which means mono-phyly of an animal group can be ascertained. The shortcomings of Hennig’s methodology are overcome and rectified in the constructivistic approach to phylogeny advocated herein. Furthermore the indispensible aspects of Hennig’s methodology are incorporated in a more general concept of phylogenetic reconstruction which was repeatedly corroborated by attempts to trace the transformation series of several fossil and recent groups of organisms.     

Hennig's enkaptic system

Hennig's phylogenetic system is characterized by an asymmetry between (sexually reproducing) species that form tokogenetic systems, versus monophyletic taxa that form a phylogenetic system. This was claimed to reflect a conflict between two hierarchies, i.e. the hierarchy of species-lineages splitting and splitting again as opposed to the phylogenetic hierarchy of groups within groups. Some cladists have sought the unification of the phylogenetic system by abandoning the species concept. In contrast, contemporary commentators (Klaus Günther, Walter Zimmermann) characterized Hennig's system as an enkaptic hierarchy. This paper explores the concept of enkapsis, and the way Hennig used it as a basis for the unification of his phylogenetic system.
 © The Willi Hennig Society 2009.     

From types to individuals: Hennig’s ontology and the development of phylogenetic systematics

Contemporary phylogenetic systematics was framed, in part, as a response to a resurgent idealistic morphology in the German‐speaking world in the first half of the 20th century. There were also conceptual and methodological challenges from Anglo‐American researchers who were sceptical about whether a phylogenetic approach to systematics could be made to work. This paper describes these challenges as a way of providing context for some ontological innovations made first by Walter Zimmermann and then by Willi Hennig. The principal argument of this paper is that what has become known as the individuality thesis played a much more important role in the conceptual foundations of Hennig’s version of phylogenetic systematics than has been widely appreciated. Understanding Hennig’s ontology illuminates his responses to objections to phylogenetic systematics from both sides of the Atlantic and sheds substantial light on the extinction part of the dichotomy rule. Although many have taken Hennig’s claim that parent species go extinct at speciation to be an arbitrary and biologically unrealistic rule, extinction of the parent follows directly from the way Hennig understands species and how they are individuated. © The Willi Hennig Society 2011.     

Grundsätze der phylogenetischen Systematik (Eine praxisorientierte Übersicht)

The essential elements of phylogenetic systematics in the sense of Hennig are emphasized: The search for synapomorphies based on a special method of comparative morphology, and the aim of an exclusive use of synapomorphies for kinship proof and the basis of systematics. Special aspects of comparative morphology are: “Directed comparisons” steady reciprocal reflection between comparative morphological result and the system methodical efforts for the realization of distinctive details, comprehensive documentation and functional interpretation. This is equally true for recent and fossil forms. Most suitable for the method (in the sense defined above) are groups with numerous differentiated morphological characters, which can also be preserved in the fossil state. The less this is the case the less is the chance for achieving necessary numbers of well proven synapomorphies. Even so, it is not permitted—for those who want to perform phylogenetic systematics in the sense of Hennig—to use convergences, parallelisms or symplesiomorphies in the sense of “synapomorphies” as phylogenetic arguments for kinship relations. Numerous examples and diagrams demonstrate the methodological proceeding, and differences towards other methods of phylogenetical reconstruction and interpretation. Special attention is paid to direct and indirect conclusions drawn from fossils: Time of origin of characters, stem groups and *groups; predictions concerning the appearance (set of characters) of fossils and simultaneous existence of “neighbour groups” (sister groups, and more distantly related taxa).     

Willi Hennig’s dichotomization of nature

Two formal assumptions implied in Willi Hennig’s “phylogenetic systematics” were repeatedly criticized for not being biologically grounded. The first is that speciation is always dichotomous; the second is that the stem‐species always goes extinct when its lineage splits into two daughter species. This paper traces the theoretical roots of Hennig’s “principle of dichotomy”. While often considered merely a methodological principle, Hennig’s realist perspective required him to ground the “principle of dichotomy” ontologically in speciation. As a methodological principle, the adherence to a strictly dichotomously structured phylogenetic system allowed Hennig to be unequivocal in character analysis and precise in the rendition of phylogenetic relationships. The ontological grounding of the “principle of dichotomy” in speciation remains controversial, however. This has implications for the application of techniques of phylogeny reconstruction to populations of bisexually reproducing organisms (phylogeography). Beyond that, the “principle of dichotomy” has triggered an intensive debate with respect to phylogeny reconstruction at the prokaryote level. © The Willi Hennig Society 2010.     

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.     

Adolf Naef (1883–1949), systematic morphology and phylogenetics

Several authors have highlighted methodological similarities between Naef’s systematic morphology and Hennig’s phylogenetic systematics. Whereas this may indicate an influence of Naef on Hennig, the relevant issues – such as the principle of generality in character analysis and the threefold parallelism of classification, ontogeny and the Fossil Record – reach back beyond Naef and Hennig and were widely discussed in the German systematics literature of the late 19th and early 20th Century. The same is true of conceptual issues, such as the discussion of the principle of monophyly, which was first introduced by Haeckel in 1866 ( Rieppel 2011b , J Zool Syst Evol Res 49 :1). In spite of methodological and conceptual agreements, Naef’s systematic morphology differed fundamentally from Hennig’s phylogenetic systematics. Naef emphasized the role of unbiased observation and the immediate acquaintance of the investigator with the phenomena given in nature as the basis of natural science in general, and of his hierarchy of types in particular. From the hierarchy of types, Naef derived through conceptual‐logical analysis the natural system, which above the species level forms a nested hierarchy of intensionally defined classes, denoted by general names. The historical‐causal interpretation of the hierarchy of types in turn offers insight into the hypothetical reality of phylogeny. Hennig in contrast denied the possibility of theory‐free observation, indeed of assumption‐free science in general, and on that basis put metaphysical issues above epistemology. Tying individuality to spatiotemporal location, historicity and causality, Hennig took not only species (as did Naef) but also supraspecific monophyletic taxa as individuals, denoted by proper names. From the species up, the phylogenetic system thus becomes a nested hierarchy of complex wholes of increasing degrees of complexity. Diagnostic characters of species or higher taxa can then no longer define classes (as in Naef’s natural system) but are thought to indirectly indicate the phylogenetic relations on which alone the phylogenetic system is to be based.     

中国瓦娄蜗牛属系统发育关系和动物地理学初探(肺螺亚纲,柄眼目,瓦娄蜗牛科)

运用支序分析方法和Hennig86软件探讨了柄眼目瓦娄蜗牛科瓦娄蜗牛属中国种类的系统发育关系。中国瓦娄蜗牛属种类现已知16种,主要分布于中国北部,也分布在亚洲北部和西部,欧洲大部以及北美东南部。选择Acan-thinual属的代表种A．aculeata作为外群。系统分析所用特征为瓦娄蜗牛届的19个形态学特征。运用命令mh^*;和bb^*;对原始数据集进行运算,并进行逐步加权处理,结果共得到步长＝102,CI值＝61,RI值＝83的同等简约树一株。选择该树作为支序分析的首选支序图。支序分析的结果初步将中国瓦娄蜗牛属的16个种分为两群：西部瓦娄蜗牛群和东部瓦娄蜗牛群。运用扩散-隔离分析方法对该属的动物地理学进行了分析和讨论,认为瓦娄蜗牛居中国种类现有分布格局的形成可能是在中新世以前主要由该属的祖先在其起源地内蒙古一带逐步向现有分布地扩散的结果。     

Post-Haeckelian comparative biology — Adolf Naef’s idealistic morphology

Summary The present study describes the conceptual framework of Adolf Naef’s idealistic morphology as presented at the onset of the 20^th century. According to Naef, Haeckel’s and Gegenbaur’s approaches towards a phylogenetic biology were insufficient. He made it clear that Haeckel’s ideas were based on typological morphology. Thus, Haeckel’s views on comparative biology pointed back to pre-Darwinian concepts. Naef’s consequence was not to work out his own evolutionary morphology but to systematize the earlier typological concept. Consequently, he separated comparative morphology from phylogenetic studies. This idea was adopted by Hennig and was even imported into modern cladism.     

A cladistic analysis of the cockroach wasps based on morphological data (Hymenoptera: Ampulicidae)

The Ampulicidae are one of the most basal groups within the apoid wasps, the paraphyletic assemblage of wasps previously known as Sphecidae. The morphology and taxonomy of the Ampulicidae are poorly studied, and there is no strict cladistic approach on their phylogeny yet. Here we assemble morphological characters of phylogenetic significance and submit them to parsimony analyses using modern cladistic methods. This produces a well-resolved and firmly supported phylogenetic hypothesis on the generic relationships within the group. Based on our phylogenetic results a revised classification is provided, subdividing the Ampulicidae into two monophyletic subfamilies, Ampulicinae ( Ampulex and Trirogma ) and Dolichurinae, the latter comprising two monophyletic tribes, Dolichurini ( Dolichurus and Paradolichurus ) and Aphelotomini, new tribe ( Aphelotoma and Riekefella ). Based on the resulting cladogram, the historical biogeography and the fossil record of Ampulicidae are briefly discussed.
© The Willi Hennig Society 2009.     

Plethodontid salamander mitochondrial genomics: A parsimony evaluation of character conflict and implications for historical biogeography

A new parsimony analysis of 27 complete mitochondrial genomic sequences is conducted to investigate the phylogenetic relationships of plethodontid salamanders. This analysis focuses on the amount of character conflict between phylogenetic trees recovered from newly conducted parsimony searches and the Bayesian and maximum likelihood topology reported by Mueller et al. (2004 ; PNAS, 101, 13820–13825). Strong support for Hemidactylium as the sister taxon to all other plethodontids is recovered from parsimony analyses. Plotting area relationships on the most parsimonious phylogenetic tree suggests that eastern North America is the origin of the family Plethodontidae supporting the “Out of Appalachia” hypothesis. A new taxonomy that recognizes clades recovered from phylogenetic analyses is proposed. © The Willi Hennig Society 2005.     

The Gegenbaur Transformation: a paradigm change in comparative biology

The leading experts in the development of phylogenetic systematics, Walter Zimmermann and Willi Hennig, formulated their research program in opposition to (neo-) idealistic morphology as expounded by authors such as Wilhelm Troll and Adolf Naef. Idealistic morphology was synonymous with systematic morphology for Naef, who wanted it to be strictly kept separate and independent of phylogenetics. Naef conceded, however, that the natural system researched by systematic morphology is to be causally explained by the theory of descent with modification. Naef went on to compile a dictionary that would regulate the translation of the language of systematic morphology into the language of phylogenetics. The switch from idealistic morphology to phylogenetic morphology is paradigmatically exemplified in the two editions (1859, 1870) of Carl Gegenbaur's Grundzüge der vergleichenden Anatomie. This paper traces the development of phylogenetic systematics from Gegenbaur through the work of Adolf Naef to Walter Zimmermann and Willi Hennig. Hennig added to Naef's systematic morphology the dimension of time, which required an ontological replacement: Naef's natural system, a nested hierarchy of intensionally defined sets subject to the membership relation, was replaced by Hennig's phylogenetic system, an enkaptic hierarchy subject to the part-to-whole relation.     

Die evolution der pantherkatzen modell zur überprüfung der brauchbarkeit der hennigschen prinzipien der phylogenetischen systematik für wirbeltierpaläontologische studien

The species and genus group of pantherine cats (Panthera, Uncia, Neofelis) has been studied in the fields of zoology, palaeontology and biogeography as far as allowing a total mosaic pattern of its evolution, which, though it is incomplete, is free of contradictions. Thus it is suited as a model for checking the applicability of Hennig’s phyletical concepts to pure palaeontological studies in vertebrates. By this it results that the principles of the phylogenetic systematics are unquestionably useful due to their terminological clearness, but are not sufficient alone for the reconstruction of evolutionary processes. Dichotomy or radiation in evolutionary splitting of a species should not be a question of principle.     

Zur Anwendung der Hennigschen Methode in der Wirbeltierpaläontologie

The application of Hennig’s theory on mere fossil groups is being discussed on the example of different primitive tetrapods. It is being demonstrated that a genealogical reconstruction in Hennig’s sense is biased by the possibility of great error, the reasons being: a) The insufficient fossil record and the resulting meagreness of usuable characters. b) Hennig’s theory does not offer any new ideas to solve the basic problems of phylogeny (parallelism, direction of evolution). Cladistic analysis biased by these insufficiencies should be supported by anagenetic investigations, i. e. by functional morphology. Hennig’s methodology, however, should be applied in cladistic reconstruction, since it works on logic reasonings and demands transparent documentation.     

THE CLASSIFICATION OF PROBOSCIDEA: HOW MANY CLADISTIC CLASSIFICATIONS?

Abstract— Hennig conceived a method to build a "phylogenetic system", with the stipulation that a "properly drawn phylogenetic tree must be directly translatable into the language of phylogenetic systematics". Consequently, this system could be the general reference system of biology. A review of the classificatory technical improvements, conventions and rules which have been proposed for the past twenty years together with their application to the classification of the Proboscidea, leads to the conclusion that more than one formal system can be built upon one given cladogram. As words are used more frequently for communication than diagrams, schemes or graphs, the "general reference system of biology1' remains somewhere in Utopia. The "phylogenetic system" is rather more synonymous with a cladogram than with a written classification.     

The PhyloCode, types, ranks and monophyly: a response to Pickett

A report from the First International Phylogenetic Nomenclature Meeting recently published in Cladistics conveys several misconceptions about the PhyloCode and presents an erroneous interpretation of discussions that took place at that meeting. Contrary to Pickett's assertions, the PhyloCode is designed to name clades, not paraphyletic groups; the rejection of ranks has never been a fundamental principle of phylogenetic nomenclature; and specifiers under the PhyloCode differ in several ways from types under rank‐based nomenclature. © The Willi Hennig Society 2005.     

On the difference between mono-, holo-, and paraphyletic groups: a consistent distinction of process and pattern

About 50 years ago, the German entomologist Willi Hennig presented a new approach in biological systematics that he called a phylogenetic systematics. The main difference between his approach and traditional Linnean systematics was that he distinguished two new kinds of groups that he called mono- and paraphyletic groups, and whereof he considered only monophyletic groups to be natural groups. However, almost immediately after publication of his approach in English, some biological systematists commented that his monophyletic groups rather ought to be called holophyletic groups. The comment sparked a heated debate about the definition of the concept 'monophyletic groups', but the debate never reached consensus. In this paper, I claim that the controversy does not concern the definition of the concept monophyletic groups per se , but instead conceptualization of phylogenies (i.e. dichotomously branching processes) in a general sense. I discuss the relation between mono-, holo- and paraphyletic groups, and conclude that Hennig's conceptualization of phylogenies is both inconsistent and empirically wrong, whereas Linné's instead is consistent and correct.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 217–220.     

Exploring the effects of phylogenetic uncertainty and consensus trees on stratigraphic consistency scores: a new program and a standardized method

The stratigraphic record of first appearances provides an independent source of data for evaluating and comparing phylogenetic hypotheses that include taxa with fossil histories. However, no standardized method exists for calculating these metrics for polytomous phylogenies, restricting their applicability. Previously proposed methods insufficiently deal with this problem because they skew or restrict the resulting scores. To resolve this issue, we propose a standardized method for treating polytomies when calculating these metrics: the Comprehensive Polytomy approach (ComPoly). This approach accurately describes how phylogenetic uncertainty, indicated by polytomies, affects stratigraphic consistency scores. We also present a new program suite (Assistance with Stratigraphic Consistency Calculations) that incorporates the ComPoly approach and simplifies the calculation of absolute temporal stratigraphic consistency metrics. This study also demonstrates that stratigraphic consistency scores calculated from strict consensus trees can be overly inclusive and those calculated from less‐than‐strict consensus trees inaccurately describe the phylogenetic signal present in the source most‐parsimonious trees (MPTs). Therefore, stratigraphic consistency scores should be calculated directly from the source MPTs whenever possible to ensure their accuracy. Finally, we offer recommendations for standardizing comparisons between molecular divergence dates and the stratigraphic record of first appearances, a promising new application of these methods. © The Willi Hennig Society 2010.     

The genera of Diopsidae (Insecta:Diptera)

Taxonomic characters which have been used in defining the genera of Diopsidae are discussed with special reference to the structure of the head and eyestalks. In consequence, the subdivisions of this family put forward by Hennig in 1965 are revised and extended, a new scheme of three subfamilies being proposed and a possible phylogenetic sequence of genera suggested.
A key for the identification of these subfamilies and current generic taxa is given.