Evolutionary theory and systematics: relationships between process and patterns

The relevance of the Modern Evolutionary Synthesis to the foundations of taxonomy (the construction of groups, both taxa and phyla) is reexamined. The nondimensional biological species concept, and not the multidimensional, taxonomic, species notion which is based on it, represents a culmination of an evolutionary understanding. It demonstrates how established evolutionary mechanisms acting on populations of sexually reproducing organisms provide the testable ontological basis of the species category. We question the ontology and epistemology of the phylogenetic or evolutionary species concept, and find it to be a fundamentally untenable one. We argue that at best, the phylogenetic species is a taxonomic species notion which is not a theoretical concept, and therefore should not serve as foundation for taxonomic theory in general, phylogenetics, and macroevolutionary reconstruction in particular. Although both evolutionary systematists and cladists are phylogeneticists, the reconstruction of the history of life is fundamentally different in these two approaches. We maintain that all method, including taxonomic ones, must fall out of well corroborated theory. In the case of taxonomic methodology the theoretical base must be evolutionary. The axiomatic assumptions that all phena, living and fossil, must be holophyletic taxa (species, and above), resulting from splitting events, and subsequently that evaluation of evolutionary change must be based on a taxic perspective codified by the Hennig ian taxonomic species notion, are not testable premises. We discuss the relationship between some biologically, and therefore taxonomically, significant patterns in nature, and the process dependence of these patterns. Process-free establishment of deductively tested “genealogies” is a contradiction in terms; it is impossible to “recover” phylogenetic patterns without the investment of causal and processual explanations of characters to establish well tested taxonomic properties of these (such as homologies, apomorphies, synapomorphies, or transformation series). Phylogenies of either characters or of taxa are historical-narrative explanations (H-N Es), based on both inductively formulated hypotheses and tested against objective, empirical evidence. We further discuss why construction of a “genealogy”, the alleged framework for “evolutionary reconstruction”, based on a taxic, cladistic outgroup comparison and a posteriori weighting of characters is circular. We define how the procedure called null-group comparison leads to the noncircular testing of the taxonomic properties of characters against which the group phylogenies must be tested. This is the only valid rooting procedure for either character or taxon evolution. While the Hennig -principle is obviously a sound deduction from the theory of descent, cladistic reconstruction of evolutionary history itself lacks a valid methodology for testing transformation hypotheses of both characters and species. We discuss why the paleontological method is part of comparative biology with a critical time dimension ana why we believe that an “ontogenetic method” is not valid. In our view, a merger of exclusive (causal and interactive, but best described as levels of organization) and inclusive (classificatory) hierarchies has not been accomplished by a taxic scheme of evolution advocated by some. Transformational change by its very nature is not classifiable in an inclusive hierarchy, and therefore no classification can fully reflect the causal and interactive chains of events constituting phylogeny, without ignoring and contradicting large areas of corroborated evolutionary theory. Attempts to equate progressive evolutionary change with taxic schemes by Haeckel were fundamentally flawed. His ideas found 19th century expression in a taxic perception of the evolutionary process (“phylogenesis”), a merger of typology, hierarchic and taxic notions of progress, all rooted in an ontogenetic view of phylogeny. The modern schemes of genealogical hierarchies, based on punctuation and a notion of “species” individuality, have yet to demonstrate that they hold promise beyond the Haeckel ian view of progressive evolution.     

Restorative and regenerative: Exploring the concepts in the circular economy

The most recognized definition of the circular economy is that it is a restorative and regenerative economy. Despite the wide use and importance attributed to the concepts of “restoration” and “regeneration,” they are rarely defined or explained in the circular economy literature. In this context, this study critically examines the two terms, while providing guidance on their future utilization and development. Specifically, the study investigates the origin of the concepts, their adoption in frameworks that anticipated the idea of the circular economy, and their connotations in the circular economy literature. The examination supports the need for clear and distinct definitions, combined with precision in usage. From a review of the literature, restoration is a better‐defined concept than regeneration, although it needs conceptual re‐enforcement relative to the biological/ecological aspects of the circular economy. This study suggests looking in the direction of restoration ecology, a well‐established branch of ecological research. Conversely, regeneration is a symbolic/evocative term with little practical application in the context of circular systems except in the case of certain agricultural practices. Until new conceptual developments intervene, regeneration does not seem to be applicable to the economy as a whole and because of this, might be abandoned as a guiding principle of the circular economy. Unlike regeneration, restoration can be considered a core principle because it has widespread application and can be a point of reference for circular applications. This does not preclude the possibility that other concepts may be needed to augment restoration.     

How reticulated are species?

Many groups of closely related species have reticulate phylogenies. Recent genomic analyses are showing this in many insects and vertebrates, as well as in microbes and plants. In microbes, lateral gene transfer is the dominant process that spoils strictly tree‐like phylogenies, but in multicellular eukaryotes hybridization and introgression among related species is probably more important. Because many species, including the ancestors of ancient major lineages, seem to evolve rapidly in adaptive radiations, some sexual compatibility may exist among them. Introgression and reticulation can thereby affect all parts of the tree of life, not just the recent species at the tips. Our understanding of adaptive evolution, speciation, phylogenetics, and comparative biology must adapt to these mostly recent findings. Introgression has important practical implications as well, not least for the management of genetically modified organisms in pest and disease control.     

DNA cloning demonstrates high genetic heterogeneity in populations of the subaerial green alga Trentepohlia (Trentepohliales,Chlorophyta)

Mats of the green alga Trentepohlia, a genus widely distributed in the tropics as well as temperate regions, have always been perceived as homogeneous (i.e., formed by only one species). As such, their general nature and specific feature play a supportive role in the species delimitation. However, the presence of morphologically dissimilar thalli was observed under the light microscope and when cultivating a piece of a single mat. To address this, we performed DNA cloning of the rbcL gene on mat fragments of T. abietina, T. annulata, T. jolithus and T. umbrina sampled in Europe to reveal if they are composed of one or more species. We revealed that more Trentepohlia haplotypes may coexist in a single mat. In consideration of this, we conclude that the use of material isolated in unialgal culture will be almost mandatory for a taxonomic reassessment of this complicated genus. Another direct implication of this problem is that herbarium specimens consisting of field‐collected material should not be used for direct sequencing. We further hypothesize the reasons why multiple haplotypes of Trentepohlia occur more frequently in the tuft‐like mats. Possibly, fragments and/or cells of other microalgae, including other species of Trentepohlia, might be retained in such mats more easily than in the crustose trentepohlialean mats.     

有关物种概念与划分中的一些问题

关于物种的问题已经争论了几百年，但至今仍未达成完全一致，尤其是在物种的概念、物种的划分及物种的形成等主要方面的观点分歧较大，可说是学派林立、百家争鸣。引起这些分歧的根本原因是生物的极端多样性，当然，也与每个人的观察角度和认识水平不无关系。目前，最大的矛盾是，一方面大家竟相给物种想下一个简明而又高度概括的定义，另一方面在实际操作中，对不同生物却是各有其标准。这样就必然产生了很多不同的物种概念，甚至产生了混乱。此外，种下单元的划分，不论在标准还是术语方面，都存在很大问题，尤其是种和亚种的分类及系统关系问题过去和现在都被严重地忽视了。     

Species pairs of north temperate freshwater fishes: Evolution,taxonomy, and conservation

Many fish species contain morphologically, ecologically and geneticallydistinct populations that are sympatric during at least some portion oftheir life cycle. Such reproductively isolated populations act asdistinct biological species, but are identified by a common Latinbinomial. These species pairs are particularly common in freshwaterfish families such as Salmonidae, Gasterosteidae and Osmeridae and aretypically associated with postglacial lakes in north temperateenvironments. The nature of the divergences between sympatric species,factors contributing to reproductive isolation, and modes of evolutionare reviewed with particular emphasis on benthic and limnetic pairs ofthreespine sticklebacks, Gasterosteus aculeatus, and anadromous(sockeye salmon) and nonanadromous (kokanee) pairs of Oncorhynchusnerka. Phylogenetic analyses typically indicate that divergencesbetween members of replicate pairs have occurred independently and,hence, particular phenotypes are not monophyletic. Consequently,taxonomic resolution of such species complexes is a vexing problem foradherents to our traditional Linnaean classification system. Sympatricspecies pairs represent a significant component of the biodiversity oftemperate freshwater ecosystems which may be underestimated because oursystem of formal taxonomy tends to obscure diversity encompassed byspecies pairs. Conservation of such systems should be recognized as apriority without formal taxonomic designation of members of speciespairs because taxonomic resolution will likely continue to proveextremely difficult when employing traditional hierarchies andprocedures.     

A multilocus genealogical approach to phylogenetic species recognition in the model eukaryote Neurospora

To critically examine the relationship between species recognized by phylogenetic and reproductive compatibility criteria, we applied phylogenetic species recognition (PSR) to the fungus in which biological species recognition (BSR) has been most comprehensively applied, the well-studied genus Neurospora. Four independent anonymous nuclear loci were characterized and sequenced from 147 individuals that were representative of all described outbreeding species of Neurospora. We developed a consensus-tree approach that identified monophyletic genealogical groups that were concordantly supported by the majority of the loci, or were well supported by at least one locus but not contradicted by any other locus. We recognized a total of eight phylogenetic species, five of which corresponded with the five traditional biological species, and three of which were newly discovered. Not only were phylogenetic criteria superior to traditional reproductive compatibility criteria in revealing the full species diversity of Neurospora, but also significant phylogenetic subdivisions were detected within some species. Despite previous suggestions of hybridization between N. crassa and N. intermedia in nature, and the fact that several putative hybrid individuals were included in this study, no molecular evidence in support of recent interspecific gene flow or the existence of true hybrids was observed. The sequence data from the four loci were combined and used to clarify how the species discovered by PSR were related. Although species-level clades were strongly supported, the phylogenetic relationships among species remained difficult to resolve, perhaps due to conflicting signals resulting from differential lineage sorting.     

Adaptive evolutionary conservation: towards a unified concept for defining conservation units

Recent years have seen a debate over various methods that could objectively prioritize conservation value below the species level. Most prominent among these has been the evolutionarily significant unit (ESU). We reviewed ESU concepts with the aim of proposing a more unified concept that would reconcile opposing views. Like species concepts, conflicting ESU concepts are all essentially aiming to define the same thing: segments of species whose divergence can be measured or evaluated by putting differential emphasis on the role of evolutionary forces at varied temporal scales. Thus, differences between ESU concepts lie more in the criteria used to define the ESUs themselves rather than in their fundamental essence. We provide a context-based framework for delineating ESUs which circumvents much of this situation. Rather than embroil in a befuddled debate over an optimal criterion, the key to a solution is accepting that differing criteria will work more dynamically than others and can be used alone or in combination depending on the situation. These assertions constitute the impetus behind adaptive evolutionary conservation.