Frank Zachos

Natural History Museum Vienna, Austria · Mammal Collection

Evolutionary convergence in distantly related species is among the most convincing evidence of adaptive evolution. The mammalian ear, responsible for balance and hearing, is not only characterised by its spectacular evolutionary incorporation of several bones of the jaw, it also varies considerably in shape across modern mammals. Using a multivaria...

Genome‐wide technologies open up new possibilities to clarify questions on genetic structure and phylogeographic history of taxa previously studied with microsatellite loci and mitochondrial sequences. Here, we used 736 individual red deer ( Cervus elaphus ) samples genotyped at 35,701 single nucleotide polymorphism loci (SNPs) to assess the popula...

Genetic monitoring of populations currently attracts interest in the context of the Convention on Biological Diversity but needs long-term planning and investments. However, genetic diversity has been largely neglected in biodiversity monitoring, and when addressed, it is treated separately, detached from other conservation issues, such as habitat...

Efforts to monitor genetic diversity in populations vary greatly among Europeancountries. The populations and species that are most likely to experience the greatest impacts of climate change are not well covered by these efforts, which suggests an urgent need for a substantial expansion in their monitoring. Read online for free: https://rdcu.be/dv...

Taxonomic data are a scientific common. Unlike nomenclature, which has strong governance institutions, there are currently no generally accepted governance institutions for the compilation of taxonomic data into an accepted global list. This gap results in challenges for conservation, ecological research, policymaking, international trade, and othe...

When producing species classifications, taxonomists are often confronted with gray-area cases. For example, because of incipient or shallow divergence, it can be scientifically valid both to split groups of organisms into separate species and to lump them together into one species. It has been claimed that, in such cases, the ranking decision is, i...

The blue antelope or bluebuck ( Hippotragus leucophaeus ) is an extinct species of antelope that lived in South Africa until ca. 1799–1800. Disappearing only 34 years after it was described, it was the first large African mammal species to have become extinct in recent times. Therefore, current scientific knowledge of the blue antelope is limited t...

Evolutionary convergence in distantly related species is among the most convincing evidence of adaptive evolution. The mammalian ear, responsible for balance and hearing, is not only characterised by its spectacular evolutionary incorporation of several bones of the jaw, it also varies considerably in shape across modern mammals. Using a new multiv...

The Golden Eagle (Aquila chrysaetos) is widely distributed in the northern Hemisphere and is thus an ideal candidate to explore large-scale biogeographic patterns. In recent years, Golden Eagle population genetics has gained considerable attention. However, the Eastern Palearctic region, a numerical stronghold for the species, is one of the least g...

There are notoriously many different definitions of species and methods of species delimitation, forcing taxonomists to make a long range of methodological decisions in species delimitation. Because of this, there are sometimes multiple viable competing methodological paths, which could lead to different ranking (or even grouping) decisions. As a r...

Genetic monitoring of populations currently attracts interest in the context of the Convention on Biological Diversity but needs long-term planning and investments. Genetic diversity has been largely neglected in biodiversity monitoring, and when addressed is treated separately, detached from other conservation issues, such as habitat alteration du...

This handbook offers a unique collection of information on all wild mammal species living in Europe and will serve as a standard reference guide for all mammalogists and readers interested in research on mammals. This volume presents comprehensive species-specific chapters covering all aspects of ungulate biology, including palaeontology, physiolog...

The earliest known fossils of Axis are dated to the late Miocene in China. The axis deer is usually considered monotypic (Fig. 1). The monophyly of Axis (besides the chital, there are three other acknowledged species: A. calamianensis, A. kuhlii, and A. porcinus) has been corroborated by recent molecular and morphological phylogenetic studies. Ther...

Mitochondrial DNA (mtDNA) lineages are recognized as important components of intra- and interspecific biodiversity, and allow to reveal colonization routes and phylogeographic structure of many taxa. Among these is the genus Cervus that is widely distributed across the Holarctic. We obtained sequences of complete mitochondrial genomes from 13 Cervu...

An up-to-date synthesis of the biology, ecology, behaviour and conservation status of the red deer. After introducing the taxonomic status and the the systematic of the species, we provide an account of its current distribution. We then describe the main morphological, physiological and genetic features; the main life history traits (growth, surviv...

Antlers are the most conspicuous trait of cervids and have been used in the past to establish a classification of their fossil and living representatives. Since the availability of molecular data, morphological characters have generally become less important for phylogenetic reconstructions. In recent years, however, the appreciation of morphologic...

The present phylogeographic pattern of red deer in Eurasia is not only a result of the contraction of their distribution range into glacial refugia and postglacial expansion, but probably also an effect of replacement of some red deer s.l. mtDNA lineages by others during the last 50 000 years. To better recognize this process, we analysed 501 seque...

A consensus among biologists has been growing in recent years for the development of a global list of accepted species (and other taxa). To date, much discussion has focused on visions for how such a list would benefit many scientific and societal disciplines. Less emphasis has been placed on understanding the many technical challenges of compiling...

Species lists are widely used in legislation and regulation to manage and conserve biodiversity. In this paper, we explore the issues caused by the lack of an adequately governed and universally accepted list of the world's species. These include lack of quality control, duplicated effort, conflicts of interest, lack of currency, and confusion in t...

Despite the important role that species names play in multiple fields, there is no globally complete list of known and described species. This lack is a result simultaneously of the complexity of planetary biodiversity, the long history of naming species in publications from all over the world, the small number of taxonomists working on many import...

Governance is the act of governing or organizing, that is a system of rules, norms, or shared strategies to guide or regulate the actions of the governed. Since the initial development of Linnaean taxonomy, a diversity of approaches have been adopted for critical taxonomic decisions, introducing pluralism to taxonomic principles and resulting in di...

Taxonomy—the delimitation, naming, classification and documentation of species and other taxa—is an often-misunderstood discipline. Complex and at times contested, taxonomy occupies a sometimes discomforting intermediate position on a continuum from descriptive to hypothetico-deductive science. Two aspects of taxonomy that are striking to many obse...

A global consensus list of the world’s species must be based on the best available taxonomic research, and its contents should not be biased towards certain political or social aims. At the same time, users of any global list must be involved or consulted in its establishment to ensure that the list meets their needs. This paper argues that while t...

Native to southern Africa, the blue antelope (Hippotragus leucophaeus) is the only large African mammal species known to have become extinct in historical times. However, it was poorly documented prior to its extinction ~ 1800 AD, and many of the small number of museum specimens attributed to it are taxonomically contentious. This places limitation...

Information about analyzed specimens - Blue antelopes (Hippotragus leucophaeus)

Lists of species underpin many fields of human endeavour, but there are currently no universally accepted principles for deciding which biological species should be accepted when there are alternative taxonomic treatments (and, by extension, which scientific names should be applied to those species). As improvements in information technology make i...

Red deer (Cervus elaphus) throughout central Europe are influenced by different anthropogenic activities including habitat fragmentation, selective hunting and translocations. This has substantial impacts on genetic diversity and the long-term conservation of local populations of this species. Here we use genetic samples from 480 red deer individua...

This introductory volume provides an overview about the history and current status of European mammals, as well as management strategies. The remaining volumes cover comprehensive overviews of each species’ biology including paleontology, physiology, genetics, reproduction and development, ecology, habitat, diet, mortality and age determination. Th...

In a recently published paper, Taylor and colleagues discussed different approaches and interpretations of mammalian taxonomy and their bearing on more general issues such as conservation and evolutionary biology. We fully endorse the fundamental importance of taxonomy and its being grounded on scientific principles. However, we also deplore a lack...

Connectivity between golden eagle (Aquila chrysaetos) populations is poorly understood. Field studies exploring natal dispersal suggest that this raptor is a philopatric species, but with the ability to roam far. However, little is known about the population structure of the species in Europe. Our study is based on 14 microsatellite loci and is com...

Connectivity between golden eagle (Aquila chrysaetos) populations is poorly understood. Field studies exploring natal dispersal suggest that this raptor is a philopatric species, but with the ability to roam far. However, little is known about the population structure of the species in Europe. Our study is based on 14 microsatellite loci and is com...

Conservation genetics is a well‐established scientific field. However, limited information transfer between science and practice continues to hamper successful implementation of scientific knowledge in conservation practice and management. To mitigate this challenge, we have established a conservation genetics community, which entails an internatio...

The cover image is based on the Original Article Witnessing extinction: Population genetics of the last European Rollers (Coracias garrulus) in Austria and a first phylogeographic analysis of the species across its distribution range, by Carina Nebel et al., https://doi.org/10.1111/jzs.12256. Cover image © Mag. Michael Tiefenbach.

The Northern Goshawk Accipiter gentilis is a medium-sized bird of prey inhabiting boreal and temperate forests. It has a Holarctic distribution with 10 recognized subspecies. Traditionally, it has been placed within the Accipiter [gentilis] superspecies, together with Henst's Goshawk A. henstii, the Black Sparrowhawk A. melanoleucus, and Meyer's Go...

Photographs of articulated bony pelves of representatives of all major mammalian (marsupial and placental) groups. Sexual dimorphism (or the lack thereof) can also be observed by comparing male and female specimens.

Objectives The narrow human birth canal evolved in response to multiple opposing selective forces on the pelvis. These factors cannot be sufficiently disentangled in humans because of the limited range of relevant variation. Here, we outline a comparative strategy to study the evolution of human childbirth and to test existing hypotheses in primate...

Appendix S1: Supporting information Figures

Table S2 Supporting information Table S2. The number of species by higher‐order taxon used to produce each Figure in the main text, including the data source.

Table S1 Supporting information Table S1. Data on neonatal and female body mass for 284 mammalian species. Data for 266 species were taken from Tague (2016), and data for an additional twelve species (highlighted in yellow) were collated from the indicated sources. When no reference is given, data are from Tague (2016).

This introductory volume provides an overview about the history and current status of European mammals, as well as management strategies. The remaining volumes cover comprehensive overviews of each species’ biology including paleontology, physiology, genetics, reproduction and development, ecology, habitat, diet, mortality and age determination. Th...

Due to broad-scale habitat loss, European Rollers (Coracias garrulus) have been decreasing in numbers rapidly during the 20th century in parts of their European distribution range. In Austria, as of 2017, only a completely isolated relict population of two breeding pairs and a few non-breeders remained in Styria compared to about 270 pairs in the 1...

The narrow human birth canal evolved in response to multiple opposing selective forces on the pelvis. These factors cannot be sufficiently disentangled in humans because of the limited range of relevant variation. We outline a comparative strategy to study the evolution of human childbirth and to test existing hypotheses in primates and other mamma...

The species problem, despite decades of heated debates, has not been resolved yet. Recently, two new species concepts have been published, the mitonuclear compatibility species concept and the inclusive species concept. I briefly discuss them, together with a recent attempt at standardizing taxonomic decisions, in the broader framework of what I be...

In a comment published in this issue, Spartaco Gippoliti (2019) presents a critique of a recent perspective paper of mine (Zachos, 2018a). Specifically, he accuses me of diminishing taxonomy and of denying it its proper role in conservation biology. Here I respond to these criticisms and argue that taxonomy is necessary but not sufficient for delin...

Mammalian taxonomy based on the diagnosability version of the Phylogenetic Species Concept has recently been declared the only sound approach to mammalian classification. In this article, based on the underlying evolutionary ontology of species taxa, I explain the fallacy of these arguments and hold that in a ‘grey area’ after lineage sundering, co...

The cheetah (Acinonyx jubatus) consists of five subspecies, of which four are distributed over Africa, but only one critically endangered population (A. j. venaticus) exists in Asia and is present in Iran. Due to short divergence times between some of the subspecies, there have been discussions launched by the IUCN Cat Specialist Group recently to...

A corarection to this Article has been published and is linked from the HTML version of this paper. The error has not been fixed in the paper.

Game species like the red deer have been subjected to anthropogenic impacts for centuries. Translocations are often carried out—sometimes illegally—not only for sporting purposes, but also to increase trophy quality, reduce inbreeding, or mitigate bottlenecks after excessive persecution. Apart from the blurring of large-scale genetic structure, tra...

We present ancient mitochondrial DNA analyses of 31 complete cytochrome b gene sequences from subfossil red deer remains from the Tyrrhenian islands (Corsica and Sardinia) and mainland Italy in a European-wide phylogeographic framework. Tyrrhenian and North African red deer, both going back to human introductions, were previously the only red deer...

DNA-barcoding is a rapidly developing method for efficiently identifying samples to species level by means of short standard DNA sequences. However, reliable species assignment requires the availability of a comprehensive DNA barcode reference library, and hence numerous initiatives aim at generating such barcode databases for particular taxa or ge...

Sequences of mitochondrial pseudogene copies integrated into the nuclear genome of stone martens (Martes foina), obtained with primer pair M_int4, and correct stone marten barcode fragments obtained with the alternative primer pair A_int4. (FAS)

Agarose gel picture showing the lack of amplification success of the entire barcoding region in stuffed and tanned hide samples. The only sample that worked was the positive control, an ethanol-preserved European hare sample (NMW 69075). For more information on samples (species, preservation method) see Table 3. (TIF)

Sequences used for designing primers. (XLSX)

As stated in the title of this chapter, this is only a very short overview. It aims at giving a readable historical summary for biologists and, more particularly, at pointing out a number of misconceptions about the historical development of the notions about species in biology. A more detailed account can be found in the first chapters of Richards...

There are about 30 species concepts in the literature. The exact number depends on what one considers a full or acknowledged concept (sometimes a comment or definition in the literature may not be called a concept and/or may not have made it into the canon of “official” species concepts), and it also depends on where one draws the line between simi...

Species concepts, or more generally, the species problem, are among the most debated issues in biology. Answers to the questions of what a species is, in what ways species really exist (if in fact they do) and how species can not only be defined but also recognized and delimited, belong at least as much, and some of them rather more, to the realm o...

Perhaps the single most important message of this book is that boundaries in nature are fuzzy. The Tree of Life is a fractal pattern with lineages nested within lineages. “Carving up nature at its joints” is therefore a messy business. Also, many phenomena relevant to species definition and delimitation (such as interbreeding, sexuality, similarity...

The introductory quotation by Aristotle is not only a nice encapsulation of the idea of a great chain of being or , but it also shows that, from the birth of Western science and philosophy, it has always been obvious that nature has fuzzy boundaries and that demarcation of similar entities is a difficult task. In Sect. 5. 2 a potential theoretical...

This chapter is best viewed as an attempt at going a little bit deeper into various aspects of the species problem while at the same time serving the overall aim of this book, namely, to provide biologists with a distillation, as it were, of the species debate(s). Therefore, none of the subchapters claim to present an exhaustive discussion of its t...

So far we have mainly dealt with the species problem from an academic perspective—how species notions have evolved historically, what ontological or metaphysical status species have, what species concepts there are and how they differ and why, etc. It was only in the last chapter on species delimitation that taxonomic practice was given more attent...

I have already stated that the species problem is in large parts a philosophical rather than a biological issue. For no other part of the problem is this as true as for the question of the ontological status of species, i.e. the question “what kind of things” species taxa are. The usual suspects for the answer to this are classes, natural kinds (wh...

Effective population size (Ne) is one of the most important parameters in population genetics and conservation biology. It translates census sizes of real populations into the size of an idealized population showing the same rate of loss of genetic diversity as the real populations under study. Several conceptually different types of Ne can be dist...

Anthropogenic fragmentation and management practices restrict the distribution and dispersal of red deer (Cervus elaphus) throughout Germany. This can lead to a reduced exchange of individuals and therefore limit gene flow between populations. For example, previous studies found very low levels of genetic diversity and strong signs of inbreeding fo...

The IBA R&C-Grant supports an on-going genetic study assessing the function of the wildlife corridor around the town Baile Tusnad for brown bears (Ursus arctos) and investigating the existence of matrilinear assemblages of food-conditioned individuals in human-bear conflict areas. In the first step, we collected DNA samples from April until Decembe...

European populations of the white-tailed eagle (Haliaeetus albicilla) suffered a drastic decline during the 20th century. In many countries, only a few dozen breeding pairs survived or the species disappeared completely. By today, the populations have recovered, naturally or through restocking (e.g. in Scotland or the Czech Republic). In the Carpat...

We analysed more than 600 red deer (Cervus elaphus) from large parts of its European distribution range at 13 microsatellite loci, presenting the first continent-wide study of this species using nuclear markers. Populations were clearly differentiated (overall FST = 0.166, Jost’s Dest = 0.385), and the BAPS clustering algorithm yielded mainly geogr...

Frank E. Zachos offers a comprehensive review of one of today’s most important and contentious issues in biology: the species problem. After setting the stage with key background information on the topic, the book provides a brief history of species concepts from antiquity to the Modern Synthesis, followed by a discussion of the ontological status...

DNA-Barcoding, die standardisierte genetische Analyse eines bestimmten Abschnitts aus dem Erbgut, ermöglicht eine rasche und kosteneffiziente Charakterisierung der Biodiversität mit einer Vielzahl von Anwendungen auch im Bereich von angewandter Forschung und Naturschutz. Ebenso wie im Naturschutz kommt Wirbeltieren als den in der Öffentlichkeit bek...

By tradition, phylogenetic trees are presented in such a way that species-poor taxa are placed on the left side and their more diverse sister taxa on the right (usually with humans on the far right). This often leads to reading the tree as a “ladder of progress” from left (allegedly primitive, basal or even ancestral) to right (allegedly advanced,...

The Golden eagle (Aquila chrysaetos) is among the most widespread birds of prey, covering basically the whole Palaearctic from Europe and North-Africa through Asia until Japan, to the North-American continent. Only few studies have addressed the species‘ genetic structure and the consequences of its demographic history so far, and none of them have...

Population genetic research has transformed from the study of genetic structures into a much wider and highly multidisciplinary research field. In this contribution, we outline the limitations of classic population genetic data and highlight the potential of combining molecular data with additional environmental, ecological and biological data sets...

All of these species have recently been split into two or more distinct species. This trend is due mainly to a shift in paradigm from the Biological Species Concept to the Phylogenetic Species Concept (PSC). While splitting may be warranted in many cases, this article argues that the PSC is a poor guide to species delimitation, particularly in its...