Trajectories to extinction: spatial dynamics of the contraction of geographical ranges

Aim We examined the range contraction of 309 declining species of animals and plants to determine if the contraction dynamics better matched predictions based on the demographic characteristics of historical populations (demographic hypothesis) or based on the contagion‐like spread of extinction forces (contagion hypothesis). Location Species included in the analysis came from all biogeographic regions. Methods We obtained range maps for 309 species from literature or through personal correspondence with authorities. Hypotheses were contrasted by examining the sequence of changes in the proportion (C) of the remnant range that fell within the central region of the historical range. Monte Carlo simulations and polynomial regressions were employed to examine changes in C during the process of range contraction. Results The results of the Monte Carlo simulations indicated that more species had observed range contractions consistent with the contagion hypothesis than expected by chance (z‐score = 2.922, P = 0.002). The Monte Carlo analysis also indicated that the number of species whose observed range contractions were consistent with the demographic hypothesis was no greater than expected by chance (z‐score = 0.337, P = 0.367). The results of the polynomial regression analysis for the two most common taxonomic groups (mammals and birds) and for all geographical regions (Australia, Africa, Eurasia, and North America) we examined also supported the contagion hypothesis. Main conclusions Most of the examined range contractions are consistent with the contagion hypothesis and that the most likely contagion is human related disturbance. These results have important implications for the conservation of endangered species.     

Geographical distribution and extinction risk: lessons from Triassic–Jurassic marine benthic organisms

Aim To evaluate the influence of geographical distribution on the extinction risk of benthic marine invertebrates using data from the fossil record, both during times of background extinction and across a mass‐extinction episode. Total geographical range is contrasted with proxies of global abundance to assess the relationships between the two essential components of geographical distribution and extinction risk. Location A global occurrence data base of fossil benthic macro‐organisms from the Triassic and Jurassic periods was used for this study. Methods Geographical distributions and biodiversity dynamics were assessed for each genus (all taxa) or species (bivalves) based on a sample‐standardized data set and palaeogeographical reconstructions. Geographical ranges were measured by the maximum great circle distance of a taxon within a stratigraphic interval. Global abundance was assessed by the number of localities at which a taxon was recorded. Widespread and rare taxa were separated using median and percentile values of the frequency distributions of occurrences. Results The frequency distribution of geographical ranges is very similar to that for modern taxa. Although no significant correlation could be established between local abundance and geographical range, proxies of global abundance are strongly correlated with geographical range. Taxon longevities are correlated with both mean geographical range and mean global abundance, but range size appears to be more critical than abundance in determining extinction risk. These results are valid when geographical distribution is treated as a trait of taxa and when assessed for individual geological stages. Main conclusions Geographical distribution is a key predictor of extinction risk of Triassic and Jurassic benthic marine invertebrates. An important exception is in the end‐Triassic mass extinction, which equally affected geographically restricted and widespread genera, as well as common and rare genera. This suggests that global diversity crises may curtail the role of geographical distribution in determining extinction risk.     

Integration of palaeontological, historical, and geographical data on the extinction of koa-finches

Identifying the root causes of extinction or endangerment requires long chronological records that begin before a population started to decline and extend until its extinction or functional extinction. We present a case study of the koa‐finches, genus Rhodacanthis, an extinct group of Hawaiian honeycreepers that was specialized to feed on green pods and seeds of the koa tree or other leguminous plants. Six island populations of koa‐finches are known; four in the Holocene fossil record and two that survived until the 1890s. We document the palaeoecological context of the fossils and identify constraints on the age span of the specimen record for each population using stratigraphic contexts, associated radiometric determinations, and museum specimen data. We estimate the potential geographical range of koa‐finches at the time of human arrival using two methods: assessment of their historical and palaeo‐habitats, and geographical information system mapping of the pre‐human distribution of the koa plant (Acacia koa) and its sister species, the koai‘a plant (Acacia koaia). After integrating the foregoing data with chronological records and distributional maps of the potential forcing agents of extinction, we conclude that at least two extinctions of island populations were due to ecological change in the lowlands in the prehistorical and perhaps the early historical periods. In the same time frame, the koa‐finch populations on Hawai‘i Island became rare and restricted to upland refugia, making them vulnerable to the upland forest harvesting and degradation that was accelerating in the 1890s. Neither climatic variation nor mosquito‐vectored diseases are likely to have caused the observed extinctions. This study illustrates an approach that can be applied to many other extinct and endangered island species to better understand the causes of high extinction rates in the human era.     

The fossil record of the sixth extinction

Comparing the magnitude of the current biodiversity crisis with those in the fossil record is difficult without an understanding of differential preservation. Integrating data from palaeontological databases with information on IUCN status, ecology and life history characteristics of contemporary mammals, we demonstrate that only a small and biased fraction of threatened species (< 9%) have a fossil record, compared with 20% of non‐threatened species. We find strong taphonomic biases related to body size and geographic range. Modern species with a fossil record tend to be large and widespread and were described in the 19^th century. The expected magnitude of the current extinction based only on species with a fossil record is about half of that of one based on all modern species; values for genera are similar. The record of ancient extinctions may be similarly biased, with many species having originated and gone extinct without leaving a tangible record.     

Ecophysiological steps of marine adaptation in extant and extinct non-avian tetrapods

Marine reptiles and mammals are phylogenetically so distant from each other that their marine adaptations are rarely compared directly. We reviewed ecophysiological features in extant non-avian marine tetrapods representing 31 marine colonizations to test whether there is a common pattern across higher taxonomic groups, such as mammals and reptiles. Marine adaptations in tetrapods can be roughly divided into aquatic and haline adaptations, each of which seems to follow a sequence of three steps. In combination, these six categories exhibit five steps of marine adaptation that apply across all clades except snakes: Step M1, incipient use of marine resources; Step M2, direct feeding in the saline sea; Step M3, water balance maintenance without terrestrial fresh water; Step M4, minimized terrestrial travel and loss of terrestrial feeding; and Step M5, loss of terrestrial thermoregulation and fur/plumage. Acquisition of viviparity is not included because there is no known case where viviparity evolved after a tetrapod lineage colonized the sea. A similar sequence is found in snakes but with the haline adaptation step (Step M3) lagging behind aquatic adaptation (haline adaptation is Step S5 in snakes), most likely because their unique method of water balance maintenance requires a supply of fresh water. The same constraint may limit the maximum body size of fully marine snakes. Steps M4 and M5 in all taxa except snakes are associated with skeletal adaptations that are mechanistically linked to relevant ecophysiological features, allowing assessment of marine adaptation steps in some fossil marine tetrapods. We identified four fossil clades containing members that reached Step M5 outside of stem whales, pinnipeds, sea cows and sea turtles, namely Eosauropterygia, Ichthyosauromorpha, Mosasauroidea, and Thalattosuchia, while five other clades reached Step M4: Saurosphargidae, Placodontia, Dinocephalosaurus, Desmostylia, and Odontochelys. Clades reaching Steps M4 and M5, both extant and extinct, appear to have higher species diversity than those only reaching Steps M1 to M3, while the total number of clades is higher for the earlier steps. This suggests that marine colonizers only diversified greatly after they minimized their use of terrestrial resources, with many lineages not reaching these advanced steps. Historical patterns suggest that a clade does not advance to Steps M4 and M5 unless these steps are reached early in the evolution of the clade. Intermediate forms before a clade reached Steps M4 and M5 tend to become extinct without leaving extant descendants or fossil evidence. This makes it difficult to reconstruct the evolutionary history of marine adaptation in many clades. Clades that reached Steps M4 and M5 tend to last longer than other marine tetrapod clades, sometimes for more than 100 million years.     

Global climate change and accuracy of prediction of species' geographical ranges: establishment success of introduced ladybirds (Coccinellidae, Chilocorus spp.) worldwide

AimPredictions of how the geographical ranges of species change implicitly assume that range can be determined without invoking climate change. The aim here was to determine how accurate predictions of range change might be before entertaining global climatic change. LocationWorldwide. MethodsAll the documented global biological control translocations of ladybirds (Coccinellidae: Chilocorus spp.) were analysed with the ecoclimatic program, CLIMEX. This program determines species distributions in relation to climate, and can be used to express the favourableness of different localities for a species. CLIMEX is also a useful exploratory tool for determining the likelihood of establishment of species introduced from one area to another. ResultsPredictive models were developed based on the likelihood of establishment of fifteen Chilocorus spp. relative to their physiological characteristics and climatic tolerances. This likelihood was compared with actual establishment with a resultant range of 0% accuracy to 100% accuracy. Only four (26.7%) species climatic tolerances could the predicted with 100% certainty. The general lack of accurate prediction was because climate is not always the overriding feature determining whether a species will establish or not. Other determinants, such as localized response to microclimate, phenology, host type and availability, presence of natural enemies and hibernation sites play a varying role over and above climate in determining whether a species will establish at a new locality. Main conclusionsThis study shows that even in the absence of climate change, range cannot always be determined, which means that most predictions of range change with climate change are likely to be wrong.     

The demonstration of speciation in fossil molluscs and living fishes

Contrary to a recent assertion, freshwater (and marine) prosobranch gastropods and freshwater bivalves are subject to considerable variability. This, and the lack of a detailed understanding of the taxonomy of the forms involved, makes it difficult to accept that the changes documented by Williamson (1981) in a fossil sequence from Lake Turkana (Africa) represent speciation events. That 10 lineages, involving gastropods and bivalves, should change simultaneously, and the deviant forms should then simultaneously become extinct, can, we believe, be more plausibly attributed to ecophenotypic responses to environmental changes than to speciation. In revealing the pattern and process of evolution, both fossil and living forms are helpful, but in demonstrating the fine-scale events during and after speciation in living animals one can utilize techniques and observations that cannot be applied to fossil material. African cichlid fishes are particularly informative in this respect. Their current explosive radiation can be interpreted as a punctuational event in evolution.     

The applicability of Rapoport's rule to the marine molluscs of the Americas

Aim We evaluated the applicability of Rapoport's rule (RR) to the marine molluscs of the Americas. The biogeographical pattern predicted by RR has been the subject of a large number of studies, some supporting it and some not. In this exercise, we attempted to generate results free of biases in taxonomy or geographical scale. Location The study area encompassed the Pacific and Atlantic sides of the entire North and South American continents. Methods Our analysis was based on secondary data. We tested the relationship of the geographical range to gradients of latitude and depth, using the methodologies of Stevens (1989) and midpoint ( Rohde et al., 1993 ). By Spearman's correlation, we related the mean amplitude of the geographical distribution to each gradient. We compared all known molluscan species together, and performed a second analysis limited to certain taxonomically well‐known groups. Results Our results were generated from a databank encompassing 4067 species. The analyses corroborated RR on both the Pacific and Atlantic coasts. In applying the pattern to the Atlantic coast, certain methodological issues had to be considered, such as the exclusion of deep‐water species and taxonomically biased groups. Regional features, such as the size of a biogeographical province, seemed to strongly affect the form of the pattern. The results also supported the association of RR with a depth gradient.     

Pattern and process in the geographical ranges of freshwater fishes

North American freshwater fishes were studied to determine whether they displayed the same relationships between log (geographical range size) and log (body size) and the same pattern of range shape as found among North American birds and mammals. The forces that produce these patterns were also investigated. The log (geographical range size) : log (body size) relationship was analysed for 121 North American freshwater fish species. Thirty‐two imperilled species were compared with 89 non‐imperilled species to determine if the overall relationship could result from differential extinction. Range geometries were analysed, within and among habitat guilds, to determine if general patterns could be detected. The log (geographical range size) : log (body size) pattern among freshwater fish species was triangular and qualitatively similar to that found for North American birds and mammals. The results suggest that below a minimum geographical range, the likelihood of extinction increases dramatically for freshwater fishes and that this minimum range size increases with body size. The pattern of fish species’ range shapes differs from that found for other North American vertebrate taxa because, on average, fish possess much smaller ranges than terrestrial species and most fish species’ geographical ranges extend further on a north–south axis than on an east–west axis. The log (geographical range size) : log (body size) pattern reveals that fish species’ geographical ranges are more constrained than those of terrestrial species. The triangular relationship may be caused by differential extinction of species with large bodies and small geographical ranges as well as higher speciation rates of small‐bodied fish. The restricted geographical ranges of freshwater fishes gives them much in common with terrestrial species on oceanic islands. Range shape patterns within habitat guilds reflect guild‐specific historical and current ecological forces. The overall pattern of range shapes emerges from the combination of ecologically different subunits.     

Species range size distributions for some marine taxa in the Atlantic Ocean. Effect of latitude and depth

The range size distributions of 6643 species in ten different fish and invertebrate taxa dwelling in pelagic (latitudinal range sizes) and benthic (latitudinal and depth range sizes) habitats on both sides of the Atlantic Ocean (80°N−70°S) were studied. The objectives were to analyse: (1) the range size distribution patterns for the various taxa and whether they have right/left skewed or lognormal distributions; (2) the geographical species distributions, to ascertain whether the distribution ranges change with latitude (Rapoport's rule); and (3) the relationship between the depth ranges of benthic species and their maximum depth of occurrence and how depth range size distributions change with latitude. The pelagic taxa exhibited larger range sizes than did the benthic taxa, continental slope/rise species excepted. On the other hand, the boundaries between geographical provinces for both benthic taxa and pelagic taxa tended to occur in association with major oceanographic processes. The shape of the latitudinal range frequency distributions (LRFDs) of the pelagic organisms were distinctly left‐skewed, and the LRFDs for most taxa were significantly different from lognormal. There was no common pattern for the distributions of the benthic organisms, which were lognormal in Cephalopoda, Stomatopoda, and Crustacea Decapoda and tended to be left‐skewed and significantly different from lognormal in Pisces. The applicability of Rapoport's rule was not clearly inferable from the results, and the rule appears to be conditioned by the location of biogeographical boundaries and the endemism rate in the different biogeographical provinces. A clear increase in depth range size with maximum depth range was observable for benthic species, confirming previous studies. Species’ depth range distributions displayed a discernible latitudinal pattern, right‐skewed at high latitudes and left‐skewed at low latitudes. The location of biogeographical boundaries, and endemism rate by biogeographical province were considered to be the factors most useful in explaining species’ distribution patterns and their conformity or nonconformity to Rapoport's rule. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80 , 437–455.     

Distribution and speciation in marine intertidal tardigrades: testing the roles of climatic and geographical isolation

Aim To analyse the importance of climatic and geographical isolation in determining the patterns of speciation and distribution of species within the tardigrade genus Echiniscoides. Location Marine intertidal zone, globally. Methods DNA was extracted from 465 individual tardigrades from 48 localities world‐wide. The tardigrades were divided into clusters using several distance‐based criteria. The phylogeny of these clusters was estimated with Bayesian analyses. The relationships between genetic distance and substrate, climate, and geographical distance were tested with a new improved Mantel test which incorporates phylogenetic uncertainties by analysing the raw tree data instead of the averaged tree. Results Approximately 40 clusters, each probably corresponding to species, were recovered from the genetic analysis; the number of clusters fluctuated depending on the criterion used for cluster delimitation. Each cluster had a limited temperature range and all clusters were confined to single oceans under all realistic criteria for cluster delimitation. Apart from a tropical cluster, each cluster occurred only in one hemisphere. Occurrence on different substrata was not correlated with genetic distance between clusters. Both geographical distance and climate were correlated with genetic distance; however, the correlation between geographical and genetic distance disappeared when the non‐independence of climatic and geographical distance were controlled for. Main conclusions The distribution of individual species of Echiniscoides is limited by climate and geographical distance. Distance does not appear to be a major factor influencing phylogeny in this genus, but ecological speciation along a temperature gradient appears to be important.     

Distribution of South African marine benthic invertebrates applied to the selection of priority conservation areas

Available data on species distributions and endemicity were compiled and examined for 11 groups of South African marine invertebrates (2533 species). For five groups species richness adhered to a well‐documented pattern, increasing from west to east, but for the other groups species richness was highest along the south coast. Endemicity was generally highest along the south coast, and lowest along the east coast. The data base was then analysed using several types of complementarity analyses, each producing a minimum set of potential reserve areas, which cumulatively represent all invertebrate species analysed. Approaches based solely on rarity, species richness and endemicity demonstrated individual biases, suggesting a need to combine all three interests. Combining the three techniques produced similar results to the individual analyses, showing conservation priorities to be highest along the east coast. Specifically, the areas of Port Elizabeth and Durban were ranked high in all analyses. Consistently, a total of 16 sites was necessary to represent all species analysed. Comparisons with similar analyses on fish and seaweeds revealed similar findings. Existing invertebrate records were shown to be biased towards centres of high sampling activity, demonstrating a need of future sampling attention in under‐represented areas.     

Further connections between the benthic marine algal floras of the northern Arabian Sea and Japan

Identifications made on some recent collections of benthic marine algae from the Sultanate of Oman, northern Arabian Sea, have revealed several new records for this region. A total of eight species, representing five species of Rhodophyceae, two species of Phaeophyceae, and a single species each of Chlorophyceae, are newly reported for Oman. Of particular interest is the fact that these species represent new records for the Indian Ocean, some of which had previously been known from Japan and environs.     

A new look at evolution in the Galápagos: evidence from the late Cenozoic marine molluscan fauna

Endemism is not as common in the marine invertebrate fauna of the Galápagos Islands region as in the adjacent terrestrial biota. Marine invertebrates in the Galápagos are largely cosmopolitan species from the Panamic, Indo-Pacific, Californian, or Peruvian faunal provinces. However, an endemic component is also present in the fauna. The observed pattern among marine invertebrate organisms can be accounted for by at least two processes: (1) genetic continuity between mainland and island populations mediated through planktonic larvae; and (2) lower rates of intrinsic evolutionary change. The evolutionary scenario standardly applied to terrestrial organisms in the Galápagos, namely, adaptive radiation and speciation in reproductive isolation from mainland source populations, does not apply to all marine invertebrates. Evidence in support of the alternative scenario for marine invertebrates comes from both published records of species occurring in the islands and recent studies of fossil-bearing deposits on several islands in the archipelago. Two misconceptions–considering the islands and sedimentary deposits to be older than now thought, and equating the rate of evolution of the terrestrial biota with the marine biota–can lead to an incorrect interpretation of evolution in the Galápagos Contrasts between marine invertebrate and terrestrial organisms serve to illustrate some fundamental differences which have important evolutionary implications. Some of these are: endemism; dispersal; taxonomic relationships; island definitions; rates of evolutionary change; and age of fossils. In terms of Darwin's evolutionary scenario, terrestrial organisms represent the paradigm and marine organisms represent the paradox.     

European butterfly populations vary in sensitivity to weather across their geographical ranges

Aim

The aim was to assess the sensitivity of butterfly population dynamics to variation in weather conditions across their geographical ranges, relative to sensitivity to density dependence, and determine whether sensitivity is greater towards latitudinal range margins.

Location

Europe.

Time period

1980–2014.

Major taxa studied

Butterflies.

Methods

We use long‐term (35 years) butterfly monitoring data from > 900 sites, ranging from Finland to Spain, grouping sites into 2° latitudinal bands. For 12 univoltine butterfly species with sufficient data from at least four bands, we construct population growth rate models that include density dependence, temperature and precipitation during distinct life‐cycle periods, defined to accommodate regional variation in phenology. We use partial R² values as indicators of butterfly population dynamics' sensitivity to weather and density dependence, and assess how these vary with latitudinal position within a species' distribution.

Results

Population growth rates appear uniformly sensitive to density dependence across species' geographical distributions, and sensitivity to density dependence is typically greater than sensitivity to weather. Sensitivity to weather is greatest towards range edges, with symmetry in northern and southern parts of the range. This pattern is not driven by variation in the magnitude of weather variability across the range, topographic heterogeneity, latitudinal range extent or phylogeny. Significant weather variables in population growth rate models appear evenly distributed across the life cycle and across temperature and precipitation, with substantial intraspecific variation across the geographical ranges in the associations between population dynamics and specific weather variables.

Main conclusions

Range‐edge populations appear more sensitive to changes in weather than those nearer the centre of species' distributions, but density dependence does not exhibit this pattern. Precipitation is as important as temperature in driving butterfly population dynamics. Intraspecific variation in the form and strength of sensitivity to weather suggests that there may be important geographical variation in populations' responses to climate change.     

Ecological correlates of island incidence and geographical range among British butterflies

The incidence of butterflies on British islands and their geographical (latitudinal) ranges are regressed on ecological and life history variables. The objective has been to investigate the contribution of individual variables and to incorporate information on phylogenetic links. The findings confirm the close relationship of species' incidence on islands with their geographical ranges on mainland Britain and that of species' geographical ranges with ecological variables, particularly migration capacity, hostplant type (variety) and breeding-habitat range. The results for island incidence considering phylogenetic links are virtually identical to those disregarding them. For geographical range, the results are similar. The key variable in each case is dispersal, scored in either one of two different ways. However, hostplant type takes precedence over breeding-habitat range when phylogenetic links are considered. Species categorized for upper and lower quartiles for geographical range form isolated clusters in the first two axes of a principal components analysis on a set of seven ecological variables. This result suggests the combined influence of a number of ecological variables on range size. Species with wide geographical ranges tend to have high migration indices, a wide variety of hostplants and ubiquitous hostplants, long flight periods and are often multi-brooded; those with narrow geographical ranges tend to have low migration indices, sparse and limited hostplant resources, short flight periods and are often univoltine. A number of life history variables are found to correlate significantly with geographical range, but account for only small amounts of variation. The lack of any association between range size and population abundance may well reflect the difficulty of obtaining adequate measures for abundance. However, we caution against expecting a strong correlation between range size and abundance.     

Evaluating the predicted extinction risk of living amphibian species with the fossil record

Bridging the gap between the fossil record and conservation biology has recently become of great interest. The enormous number of documented extinctions across different taxa can provide insights into the extinction risk of living species. However, few studies have explored this connection. We used generalised boosted modelling to analyse the impact of several traits that are assumed to influence extinction risk on the stratigraphic duration of amphibian species in the fossil record. We used this fossil‐calibrated model to predict the extinction risk for living species. We observed a high consensus between our predicted species durations and the current IUCN Red List status of living amphibian species. We also found that today's Data Deficient species are mainly predicted to experience short durations, hinting at their likely high threat status. Our study suggests that the fossil record can be a suitable tool for the evaluation of current taxa‐specific Red Listing status.