The Oxford Dodo. Part 1: the museum history of the Tradescant Dodo: ownership,displays and audience

The Dodo Raphus cucullatus, a giant flightless pigeon endemic to Mauritius, became extinct in the late seventeenth century, and so rapid was the birds’ disappearance, that by the beginning of the nineteenth century even its very existence was questioned. Only four specimens were then recorded in European museums, of which the most famous was the Tradescant or Oxford Dodo, now in the Oxford University Museum of Natural History. It comprised the head and one foot, and unique soft tissue in the form of skin and traces of feathers. The history of this specimen is reviewed, including the still unresolved question of how it came to Britain, and we provide evidence to show that it was stuffed but probably never mounted. The changes of ownership, and its cataloguing and curation in the different museums are also described, along with its varying roles in entertainment, education and research from the earliest years until the nineteenth century. This is part one of a two-part article; the second deals with the Tradescant Dodo from its dissection in the 1840s until the present day.     

Analysis of geographic centrality and genetic diversity in the declining grasshopper species Bryodemella tuberculata (Orthoptera: Oedipodinae)

Human-induced ecological and climatic changes have led to the decline and even local extinction of many formerly widely distributed temperate and cold-adapted species. Determining the exact causes of this decline remains difficult. Bryodemella tuberculata was a widely distributed orthopteran species before the mid-19th century. Since then, many European populations have suffered drastic declines and are now considered extinct or critically endangered. We used ecological niche modelling based on a large dataset of extant and extinct occurrence data to investigate whether poor climatic suitability in the periphery of its global range was a possible cause of the local extinction of the European populations of B. tuberculata. We also used population genetics based on the COI marker to estimate and compare the genetic diversity of extant populations. We found that Europe still provides highly suitable habitats close to the climatic optimum, contradicting the assumption of climate change as major driver of this decline. Instead, changes in land-cover and other anthropogenic modifications of the habitats at the local scale seem to be the major reasons for local extinctions. Genetic analysis suggests Central Asia as center of diversity with a stable population size, whereas the effective sizes of the remaining European populations are decreasing. We found European genetic lineages nested within Central Asian lineages, suggesting a Central Asian source distribution area. Our results suggest that the declining European populations represent relics of a formerly wider distribution, which was fragmented by changes in land-use. These relics are now threatened by limited connectivity and small effective population sizes. Specific conservation actions, such as the restoration of former or potential new habitats, and translocation of individuals from extant populations to these restored sites may help slow, stall, or even revert the extinction process.

     

The Oxford Dodo. Part 2: from curiosity to icon and its role in displays,education and research

The Tradescant or Oxford Dodo has played an important role in exhibition and education throughout its 360-year history, and has been the subject of scientific research reflecting changing interests and techniques over this time. Due to confusion over its relationships, its placement in the classification systems continually changed, until the dissection in 1847 of the head and foot confirmed its columbid (pigeons and doves) affinities. Here, we describe the dissection of the head and foot and the Tradescant Dodo’s display history, from the late nineteenth century until the present day, and also its use in education. We discuss the importance of the Tradescant Dodo to Lewis Carroll, whose ‘Alice’s Adventures in Wonderland’ made the dodo aware to a worldwide audience. We further provide an overview of recent work including electron microscopic study of feathers, mDNA analysis, cytological investigation of the skin and measuring of the brain capacity, all of which have added to our knowledge of the evolution and ecology of this most extraordinary of birds. Research is ongoing; the Tradescant Dodo is presently subject to CT scanning and functional analysis. This is the second part of a two-part paper.     

Dark extinction: the problem of unknown historical extinctions

The extinction of species before they are discovered and named (dark extinction, DE) is widely inferred as a significant part of species loss in the ‘pre-taxonomic’ period (approx. 1500–1800 CE) and, to some extent, in the ‘taxonomic period’ (approx. 1800–present) as well. The discovery of oceanic islands and other pristine habitats by European navigators and the consequent introduction of destructive mammals, such as rats and goats, started a process of anthropogenic extinction. Much ecosystem change happened before systematic scientific recording, so has led to DE. Statistical methods are available to robustly estimate DE in the ‘taxonomic period’. For the ‘pre-taxonomic period’, simple extrapolation can be used. The application of these techniques to world birds, for example, suggests that approximately 56 DEs occurred in the ‘taxonomic period’ (1800–present) and approximately 180 in the ‘pre-taxonomic period’ (1500–1800). Targeting collection activities in extinction hotspots, to make sure organisms are represented in collections before their extinction, is one way of reducing the number of extinct species without a physical record (providing that collection efforts do not themselves contribute to species extinction).     

Species–area relationships of butterflies in Europe and species richness forecasting

Species–area relationships (SARs) of European butterfly species (Rhopalocera) appear to follow power functions with Mediterranean butterflies having a much higher slope value (z=0.49) compared to the slope for the northern and eastern European countries (z=0.10). A simulated process of species extinction by a stepwise density dependent random elimination of species affected species–area patterns differently. For Mediterranean countries SAR slopes decreased, for other European countries slopes increased during the extinction process. Comparisons of species numbers before and after extinction with those predicted by a classical SAR approach differed widely and revealed that SARs are not able to predict future species numbers at local scales. For Mediterranean countries the classical SAR approach underestimated the number of species remaining after simulated extinction, for all other European countries SARs highly overestimated species numbers. These contrasting patterns indicate that changes in SAR patterns do not unequivocally point to changes in species diversity or community structure as assumed by current theory. On the other hand, the results strongly indicate that simplified applications of SARs for forecasting might give misimpressions about species loss and future biodiversity if the initial community structure, especially relative densities and numbers of species with restricted range size, are not taken into account.     

The problem with the species problem

When Charles Darwin convinced the scientific community that species evolve, the long-held essentialist view of each species as fixed was rejected and a clear conceptual understanding of the term was lost. For the next century, a real species problem existed that became culturally entrenched within the scientific community. Although largely solved decades ago, the species problem remains entrenched today due to a suite of factors. Most of the factors that help maintain its perceived intractability have been revealed and logically dismissed; yet this is not widely known so those factors continue to be influential. It is time to recognize this false foundation and relegate the species problem to history.     

The dodo,the deer and a 1647 voyage to Japan

More has been written about the iconic Dodo Raphus cucullatus of Mauritius than any other extinct bird, yet despite its familiarity, only a few specimens were exported from Mauritius; individual birds reached Europe alive in 1626 and 1638 and at least two survived a journey to India in 1625. There are also vague records of other exported birds. Here, we provide confirmation based on seventeenth century documents that a live Dodo was sent to Japan in 1647, the last known captive bird, and comment on the details of its long and arduous voyage.     

The role of the Weberian Reform in European Rubus research and the taxonomy of locally distributed species – which species should we describe?

After Sudre published his treatment of European Rubi in the early 20th century, Rubus taxonomy in Europe suffered from a scholastic phase and a longer period of stagnation. The so‐called ‘Weberian Reform’ initiated the necessary revival of European batology. It rests on four major pillars: 1) mapping projects over larger areas, 2) evaluation of type material, 3) visits to loci classici, and 4) evaluation of the status of species by means of their distribution areas. Subsequently, it has become widely accepted in European batology that only species with a distribution area over 50 km should be described. Although this pragmatic species concept has been useful in making a continent‐wide overview of brambles, we argue that it is lacking a scientific basis, and should thus be rejected. There are at least four distinctive problems when treating locally distributed brambles: 1) primary hybrids, 2) locally distributed stabilised apomicts, 3) intraspecific variation in species with a larger distribution range, and 4) unstabilised swarms of hybridogenic biotypes and the derivates thereof (mainly in the montane regions). When facing the problems in Rubus systematics, we argue that all independently evolving lineages should be described as species, including apomictic lineages with very small distribution ranges, both from the mountain‐dwelling glandular series and from the lowlands. Neither primary hybrids (which are not stabilised by apomixis), nor biotypes without an independent and coherent distribution area are independently evolving lineages, and should thus not be described as species. We advocate a restrained attitude when describing new species with limited distribution areas.     

Pond conservation: from science to practice

In Europe, ponds are an exceptionally numerous and widely distributed landscape feature forming a major part of the continental freshwater resource and contributing significantly to freshwater biodiversity conservation. This has been reflected by a growing scientific concern over the first few years of the twenty-first century and is evidenced by an increasing number of academic publications on pond related topics, particularly those relating to biodiversity. It is essential, however, that this expanding scientific knowledge is widely disseminated to those involved with pond management and is then rapidly translated into action. Inevitably, the task of transferring science to practice remains a significant challenge. As a first step towards meeting this challenge the European Pond Conservation Network (EPCN), at its biennial meeting in 2008 in Valencia (Spain), made this the main theme of the conference together with two special workshops further encouraging exchanges between scientists, practitioners and policy makers. The papers selected for this special issue of Hydrobiologia (from over 120 communications presented) are all from the conference. They represent a diverse collection of themes from across the continent and North Africa and present new and original insights into topics as wide ranging as: pond biodiversity; human disturbance; landscape ecology; ecological assessment and monitoring; practical management measures; ecological restoration; hydrology and climate change; invasive species and threatened species. In all cases, the papers demonstrate an overriding need for the development of a tight link between scientific knowledge and management. Furthermore, scientific advances have to be beneficial for on the ground management and, vitally, have to be disseminated, communicated and implemented into local, national and international policy. As such, national and international networks (such as the EPCN) have a central role to play and have to develop a robust information and communication strategy which will enable the dissemination of best practice materials and advice across the continent and beyond. The work contained in this volume represents a step in the right direction and will help to ensure that ponds remain a characteristic and highly visible feature of the European landscape in the twenty-first century. Guest editors: B. Oertli, R. Céréghino, A. Hull & R. Miracle Pond Conservation: From Science to Practice. 3rd Conference of the European Pond Conservation Network, Valencia, Spain, 14–16 May 2008     

The evolutionary significance of mass extinctions

Local extinctions of populations, species or groups of species in a particular area are commonly observed by biologists. There are also historical records of the total extinction of single species such as the Dodo, the Great Auk and the Tasmanian Wolf. Mass extinctions are on a much larger scale, and their study is based on the fossil record. The aims of this review are to explore the nature of mass extinctions and their evolutionary significance. The key questions are: what is mass extinction, what are the causes of mass extinctions, do mass extinctions follow a regular pattern, and how do mass extinctions affect our understanding of evolutionary processes?     

Phylogeography, genetic structure and diversity in the endangered bearded vulture (Gypaetus barbatus, L) as revealed by mitochondrial DNA

Bearded vulture populations in the Western Palearctic have experienced a severe decline during the last two centuries that has led to the near extinction of the species in Europe. In this study we analyse the sequence variation at the mitochondrial control region throughout the species range to infer its recent evolutionary history and to evaluate the current genetic status of the species. This study became possible through the extensive use of museum specimens to study populations now extinct. Phylogenetic analysis revealed the existence of two divergent mitochondrial lineages, lineage A occurring mainly in Western European populations and lineage B in African, Eastern European and Central Asian populations. The relative frequencies of haplotypes belonging to each lineage in the different populations show a steep East-West clinal distribution with maximal mixture of the two lineages in the Alps and Greece populations. A genealogical signature for population growth was found for lineage B, but not for lineage A; futhermore the Clade B haplotypes in western populations and clade A haplo-types in eastern populations are recently derived, as revealed by their peripheral location in median-joining haplotype networks. This phylogeographical pattern suggests allopatric differentiation of the two lineages in separate Mediterranean and African or Asian glacial refugia, followed by range expansion from the latter leading to two secondary contact suture zones in Central Europe and North Africa. High levels of among-population differentiation were observed, although these were not correlated with geographical distance. Due to the marked genetic structure, extinction of Central European populations in the last century re-sulted in the loss of a major portion of the genetic diversity of the species. We also found direct evidence for the effect of drift altering the genetic composition of the remnant Pyrenean population after the demographic bottleneck of the last century. Our results argue for the management of the species as a single population, given the apparent ecological exchangeability of extant stocks, and support the ongoing reintroduction of mixed ancestry birds in the Alps and planned reintroductions in Southern Spain.     

Inconsistent detection of extinction debts using different methods

The extinction debt, delayed species extinctions following landscape degradation, is a widely discussed concept. But a consensus about the prevalence of extinctions debts is hindered by a multiplicity of methods and a lack of comparisons among habitats. We applied three contrasting species–area relationship methods to test for plant community extinction debts in three habitats which had different degradation histories over the last century: calcareous grassland, heathland and woodland. These methods differ in their data requirements, with the first two using information on past and current habitat area alongside current species richness, whilst the last method also requires data on past species richness. The most data‐intensive, and hence arguably most reliable method, identified extinction debts across all habitats for specialist species, whilst the other methods did not. All methods detected an extinction debt in calcareous grassland, which had undergone the most severe degradation. We conclude that some methods failed to detect an extinction debt, particularly in habitats that have undergone moderate degradation. Data on past species numbers are required for the most reliable method; as such data are rare, extinction debts may be under‐reported.     

Potential impacts of climate change on the distributions and diversity patterns of European mammals

The Intergovernmental Panel on Climate Change (IPCC) predicts an increase in global temperatures of between 1.4°C and 5.8°C during the 21st century, as a result of elevated CO₂ levels. Using bioclimatic envelope models, we evaluate the potential impact of climate change on the distributions and species richness of 120 native terrestrial non-volant European mammals under two of IPCC’s future climatic scenarios. Assuming unlimited and no migration, respectively, our model predicts that 1% or 5–9% of European mammals risk extinction, while 32–46% or 70–78% may be severely threatened (lose > 30% of their current distribution) under the two scenarios. Under the no migration assumption endemic species were predicted to be strongly negatively affected by future climatic changes, while widely distributed species would be more mildly affected. Finally, potential mammalian species richness is predicted to become dramatically reduced in the Mediterranean region but increase towards the northeast and for higher elevations. Bioclimatic envelope models do not account for non-climatic factors such as land-use, biotic interactions, human interference, dispersal or history, and our results should therefore be seen as first approximations of the potential magnitude of future climatic changes.     

Biography of a "Feathered Pig": The California Condor Conservation Controversy

In the early 20th century, after hundreds of years of gradual decline, the California condor emerged as an object of intensive scientific study, an important conservation target, and a cultural icon of the American wilderness preservation movement. Early condor researchers generally believed that the species' survival depended upon the preservation of its wilderness habitat. However, beginning in the 1970s, a new generation of scientists argued that no amount of wilderness could prevent the condor's decline and that only intensive scientific management - including captive breeding - could save the species from certain extinction. A bitter and highly politicized battle soon developed over how to best preserve the condor. For 5years the condor was extinct in the wild; however, by the time that officials released the first captive-bred birds the condor recovery program had garnered widespread public support, even among its former critics. Today, condor advocates from the scientific and activist communities work together to manage the species and protect its habitat. The condor's story illustrates some of the tensions, problems, and successes that have accompanied the rise of conservation biology as a scientific field and environmental movement in the United States.     

Metapopulation shift and survival of woodland birds under climate change: will species be able to track?

Climate change has been widely recognized as a key factor driving changes in species distributions. In this study we use a metapopulation model, with a window of suitable climate moving polewards, to explore population shifts and survival of woodland birds under different climate change scenarios and landscape configurations. Extinction vulnerability and expansion ability are predicted for the middle spotted woodpecker Dendrocopus medius and two alternative r‐K strategies under west European climate change scenarios of 1, 2 and 4°C temperature increase per century, corresponding to isotemperature velocities of ca 2, 4 and 8 km yr^?1. The simulated northward expansion of the bird's distribution is typically in the range of only 0–3 km yr^?1, in spite of 10–20 times larger maximum dispersal distances. This is too slow to track the climate change‐driven range contraction of 4 or 8 km yr^?1 in the south resulting in metapopulation extinction. Especially K‐selected (large‐bodied) species are vulnerable in the simulations. With a temperature increase of 4°C per century bird species go extinct within 104–178 yr. We present a simple approximation formula to predict the mean time to metapopulation extinction using 1) the rate of climate change, which determines the speed of range contraction in the south, 2) the size of the distribution range, which serves as a buffer against extinction, and 3) the northward expansion velocity, determined by species traits and landscape properties. Finally, our results indicate that the northward expansion rate is not constant. It will be initially lagged suggesting that recently observed expansion rates might be underestimations of future northward expansion rates.     

Classification,nomenclature and identification of lime swallowtail butterflies: A post‐cladistic analysis (Lepidoptera: Papilionidae)

Specimens taxonomically treated in the Fauna Hawaiiensis were associated by cluster analysis, thereby reconstructing assemblages of Hawaiian carabid beetle species (Coleoptera: Carabidae) observed during the late 19th century. Associations among specimens representing 193 species permit concise hypotheses of habitat preferences for many of the 32 carabid species collected during the early period of European scientific exploration (1872–1902), but not observed since. These associations are consistent with data derived from contemporary biological surveys of Hawai'i. Absence of entire clusters of associated species from recent collections suggests actions of common agents leading to extinction or extreme population reduction. The candidate list of threatened and endangered species of the US. Fish and Wildlife Service established prior to 1994 included one Hawaiian carabid species missing since 1902, versus eight other species collected at various times over the past century. Improvements in knowledge of carabid beetle species’ spatial distribution and temporal persistence derived from recent field survey and taxonomic research demonstrate that the types of criteria used to construct that list must be rejected. Future consideration of official conservation status for any Hawaiian carabid beetle species must take into account the status of ecologically associated species, and the limited likelihood that individuals of all extant species can be consistently observed in nature due to their natural relative rarity or their secretive habits within restricted geographic and ecological distributions. Historical specimen associations serve as the best guides for continuing efforts to monitor known faunal members and to rediscover long‐missing species. These associations also serve to link information concerning individual species with particular habitats.     

Environmental and human factors influencing rare plant local occurrence, extinction and persistence: a 115-year study in the Mediterranean region

Aim Assessing whether environmental and human factors influenced the spatial distribution and the dynamics of regionally rare plant species since the late nineteenth century, and whether these spatial and temporal patterns of rare species occurrences differ according to their chorology (level of endemism and biogeographic affinity). Location An area extending over 6250 km² in the French Mediterranean Region. Methods We used two botanical surveys achieved in 1886 and in 2001, and considered species rare if occurring in only one or two sites in the study area. Each rare species was assigned to a group of endemism level (restricted endemic, non‐endemic), and of biogeographic affinity (Mediterranean, South/Central European, Mountain, Eurosiberian). A 1 × 1 km grid was applied to the study zone. Generalized linear models were developed to study the spatial distribution and the fate of rare species occurrences (local extinction vs. local persistence between 1886 and 2001), as a function of environmental and human variables. Multivariate analyses were used to test whether the spatial distribution and the fate of rare species occurrences differed according to their chorology. Results In 2001, rare species as a whole tended to occur at higher altitude, in zones dominated by semi‐natural open habitats, and where cultivated area had decreased in the last 30 years. Between 1886 and 2001, rare species were the most prone to local extinction in zones where human population density, cultivated area and livestock density had increased the most. Between 1886 and 2001, rare species had a higher probability of local persistence in zones of high altitude and steep slope, on basic bedrocks and with low cultivated area. Rare species with Mountain and Eurosiberian affinities occurred in marginal habitats in the study region, i.e. on gneiss‐micaschist bedrocks and at high altitudes, whereas Mediterranean and South/Central European rare species occupied more varied environmental conditions. Between 1886 and 2001, Eurosiberian rare species showed high rates of local extinction whereas Mediterranean rare species had a significantly higher probability of local persistence. Restricted endemic species mostly occurred in zones of high slope, low human population density, and where cultivated area had decreased in the last 30 years. Occurrences of restricted endemics remained significantly stable between 1886 and 2001. Main conclusions Environmental and land‐use changes that occurred over the twentieth century in the Mediterranean Basin had significant impacts on the spatial distribution and on the long‐term dynamics of rare species occurrences. Urbanization and recent agriculture intensification, occurring mainly in coastal plains and littoral zones, caused most local extinctions of rare species from 1886 to 2001. Local populations of Eurosiberian species, which reach their range limits in marginal zones of the Mediterranean, also appear to be highly vulnerable. Conversely, most restricted endemic species occur in habitats with harsh topography and low human disturbance and have a higher potential of local persistence.     

Viability and Management of an Endangered Capercaillie (Tetrao urogallus) Metapopulation in the Jura Mountains, Western Switzerland

The populations of Capercaillie (Tetrao urogallus), the largest European grouse, have seriously declined during the last century over most of their distribution in western and central Europe. In the Jura mountains, the relict population is now isolated and critically endangered (about 500 breeding adults). We developed a simulation software (TetrasPool) that accounts for age and spatial structure as well as stochastic processes, to perform a viability analysis and explore management scenarios for this population, capitalizing on a 24 years-long series of field data. Simulations predict a marked decline and a significant extinction risk over the next century, largely due to environmental and demographic stochasticity (average values of life-history parameters would otherwise allow stability). Variances among scenarios mainly stem from uncertainties about the shape and intensity of density dependence. Uncertainty analyses suggest to focus conservation efforts on enhancing, not only adult survival (as often advocated for long-lived species), but also recruitment. The juvenile stage matters when local populations undergo extinctions, because it ensures connectivity and recolonization. Besides limiting human perturbations, a silvicultural strategy aimed at opening forest structure should improve the quality and surface of available patches, independent of their size and localization. Such measures are to be taken urgently, if the population is to be saved.     

Testing for equal rates of cladogenesis in diverse taxa

Taxa differ widely in numbers of species, which may be due either to chance alone or to factors that cause differences in speciation and extinction rates between taxa. To test whether an observed distribution of species over taxa differs from the distribution expected from chance alone, one must take into account that neither speciation nor extinction rates are known. This paper introduces a way to estimate speciation and extinction probabilities from the distribution of extant species over families and to test whether the observed distribution is different from expected. Application of this procedure to the distributions of bird, hexapod, primate, and angiosperm species over taxa provides statistical evidence of differences in rates of cladogenesis between taxa.     

Morphological variability in time and space: an example of patterns within buchiid bivalves (Bivalvia,Buchiidae)

Abstract: We studied morphological variation of the bivalve Buchia over its geographical and temporal range. Buchia was widely distributed during the Late Jurassic and Early Cretaceous, and, while previous quantitative studies have shown that species are characterized by large amounts of variation, there have been no prior attempts to measure how morphology varies geographically. We employed traditional morphometric techniques using nine linear/angular measurements on 1855 buchiid shells from eight localities taken from widely separated, but mostly coeval, sections across its range. Principal component and canonical variate analyses indicate that geographical morphospace of buchiids varied significantly, but we did not find evidence of a latitudinal gradient in shell shape. The amount of variation between localities was similar to the amount of variation between species, indicating the importance of geographical effects on morphology. Disparity (morphological diversity within a taxon, calculated by the sum of variances) and diversity (number of species) were calculated for each location and time period (age). Disparity and diversity reached ultimate lows just before the genus’ extinction in the Hauterivian, and is suggestive that extinction was morphologically selective. We did not find significant trends for either metric, but there were discordances throughout its temporal range. Latitudinal trends of disparity and diversity within Buchia are not apparent. This research adds to the growing body of work on geographical variation and is a preliminary step to understanding the nature and variation of buchiid species and of biodiversity in general.