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Lost in time, lonely, and single: reproductive asynchrony and the Allee effect.

Identifying linkages between life-history traits and small population processes is essential to effective multispecies conservation. Reproductive asynchrony, which occurs when individuals are reproductively active for only a portion of the population-level breeding period, may provide one such link. Traditionally, reproductive asynchrony has been considered from evolutionary perspectives as an advantageous bet-hedging strategy in temporally unpredictable environments. Here, we explore the dynamic consequences of reproductive asynchrony as a density-dependent life-history trait. To examine how asynchrony affects population growth rate and extinction risk, we used a general model of reproductive timing to quantify the temporal overlap of opposite-sex individuals and to simulate population dynamics over a range of initial densities and empirical estimates of reproductive asynchrony. We also considered how protandry, a sexually selected life-history strategy that often accompanies asynchrony, modulates the population-level effects of reproductive asynchrony. We found that asynchrony decreases the number of males a female overlaps with, decreases the average probability of mating per male/female pair that does overlap, and leaves some females completely isolated in time. This loss of reproductive potential, which is exacerbated by protandry, reduces population growth rate at low density and can lead to extinction via an Allee effect. Thus reproductive asynchrony and protandry, both of which can be evolutionarily advantageous at higher population densities, may prove detrimental when population density declines.

Animals↗

A test of Simpson's "rule of the survival of the relatively unspecialized" using fossil crinoids.

Temporally long-ranging (=long-lived) taxa have been postulated to have unusual properties that aid their prolonged geologic survival. Past studies have examined dispersal capabilities, geographic ranges, and single-character morphological adaptations as factors that may contribute to geologic longevity. Here, I test whether long-lived fossil crinoid taxa are morphologically unusual using a whole suite of morphological characters. I define long-lived taxa in several explicit, comparative ways. I find that long-lived crinoid genera and families are often less distant from mean morphologies of their crinoid orders than their shorter-lived relatives; that is, they are relatively less specialized. I also compare the morphology of crinoid genera relative to basal members of their respective orders; mean morphological distances of long-lived genera from basal morphologies are seldom distinct from those of their shorter-lived relatives. I observe that long-lived crinoid genera are less distant from mean morphologies of their temporal cohorts compared with shorter-lived genera but not in a statistically significant manner. I conclude that long-lived crinoids are relatively unspecialized, in the sense that they are relatively closer to mean morphologies of their taxonomic groups.

Animals↗

Theory for designing nature reserves for single species.

We examine the question of the optimal number of reserves that should be established to maximize the persistence of a species. We assume that the mean time to extinction of a single population increases as a power of the habitat area, that there is a certain amount of habitat to be reserved, and that the aim is to determine how this habitat is most efficiently divided. The optimal configuration depends on whether the management objective is to maximize the mean time to extinction or minimize the risk of extinction. When maximizing the mean time to extinction, the optimal number of independent reserves does not depend on the amount of available habitat for the reserve system. In contrast, the risk of extinction is minimized when individual reserves are equal to the optimal patch size, making the optimal number of reserves linearly proportional to the amount of available habitat. A model that includes dispersal and correlation in the incidence of extinction demonstrates the importance of considering the relative rate at which these two factors decrease with distance between reserves. A small number of reserves is optimal when the mean time to extinction increases rapidly with habitat area or when risks of extinction are high.

Animals↗

Predator diet breadth influences the relative importance of bottom-up and top-down control of prey biomass and diversity.

We investigated the effects of predator diet breadth on the relative importance of bottom-up and top-down control of prey assemblages, using microbial food webs containing bacteria, bacterivorous protists and rotifers, and two different top predators. The experiment used a factorial design that independently manipulated productivity and the presence or absence of two top predators with different diet breadths. Predators included a "specialist" predatory ciliate Euplotes aediculatus, which was restricted to feeding on small prey, and a "generalist" predatory ciliate Stentor coeruleus, which could feed on the entire range of prey sizes. Both total prey biomass and prey diversity increased with productivity in the predator-free control and specialist predator treatments, a pattern consistent with bottom-up control, but both remained unchanged by productivity in the generalist predator treatment, a pattern consistent with top-down control. Linear food chain models adequately described responses in the generalist predator treatment, whereas food web models incorporating edible and inedible prey (which can coexist in the absence of predators) adequately described responses in the specialist predator treatment. These results suggest that predator diet breadth can play an important role in modulating the relative strength of bottom-up and top-down forces in ecological communities.

Animals↗

Host specialization and geographic localization of avian malaria parasites: a regional analysis in the Lesser Antilles.

We recovered 26 genetically distinct avian malaria parasite lineages, based on cytochrome b sequences, from a broad survey of terrestrial avifauna of the Lesser Antilles. Here we describe their distributions across host species within a regional biogeographic context. Most parasite lineages were recovered from a few closely related host species. Specialization on one host species and distribution across many hosts were both rare. Geographic patterns of parasite lineages indicated limited dispersal and frequent local extinction. The central islands of the archipelago share similar parasite lineages and patterns of infection. However, the peripheral islands harbor well-differentiated parasite communities, indicating long periods of isolation. Nonetheless, 20 of 26 parasite lineages were recovered from at least one of three other geographic regions, the Greater Antilles, North America, and South America, suggesting rapid dispersal relative to rate of differentiation. Six parasite lineages were restricted to the Lesser Antilles, primarily to endemic host species. Host differences between populations of the same parasite lineage suggest that host preference may evolve more rapidly than mitochondrial gene sequences. Taken together, distributions of avian malarial parasites reveal evidence of coevolution, host switching, extinction, and periodic recolonization events resulting in ecologically dynamic as well as evolutionarily stable patterns of infection.

Animals↗

Age structure, residents, and transients of Miocene rodent communities.

The age structures of two successive rodent communities are studied on the basis of a rich record from well-dated Miocene sections (17-10 Ma) in north-central Spain. Community age is defined as the mean of the residence times of the community members at the time of the locality age. Community ages are negatively correlated with the numbers of community members. These members are divided into residents (with continuous membership times > or =1.54 million years) and transients (with membership times <1.54 million years). During episodes of species loss, there is a preferential disappearance of transients while residents are retained, a pattern referred to as the "seniority rule." The residents define the studied communities. They are associated with early successional stages of vegetation, and transients are associated with later stages. Under stable conditions, early arrivals in succession are "transient" and replaced by competitive later arrivals. The reversed roles of transients and residents in the studied fossil record are explained by assuming high degrees of disturbance. We view the system within the context of nonequilibrium metapopulation theory, in which competitively superior species become transients because of their dependence on ephemeral late successional habitats.

Animals↗

Diversity, endemism, and age distributions in macroevolutionary sources and sinks.

Quantitative tests of historical hypotheses are necessary to advance our understanding of biogeographic patterns of species distributions, but direct tests are often hampered by incomplete fossil or historical records. Here we present an alternative approach in which we develop a dynamic model that allows us to test hypotheses about regional rates of taxon origination, extinction, and dispersal using information on ages and current distributions of taxa. With this model, we test two assumptions traditionally made in the context of identifying regions as "centers of origin"--that regions with high origination rates will have high diversity and high endemism. We find that these assumptions are not necessarily valid. We also develop expressions for the regional age distributions of extant taxa and show that these may yield better insight into regional evolutionary rates. We then apply our model to data on the biogeography and ages of extant genera of marine bivalves and conclude that diversity in polar regions predominantly reflects dispersal of taxa that evolved elsewhere rather than in situ origination-extinction dynamics.

Animals↗

Metapopulation extinction thresholds in rain forest remnants.

Although habitat fragmentation is a major threat to global biodiversity, the demographic mechanisms underlying species loss from tropical forest remnants remain largely unexplored. In particular, no studies at the landscape scale have quantified fragmentation's impacts on colonization, extinction, and local population growth simultaneously. In central Amazonia, we conducted a multiyear demographic census of 292 populations of two leaf-inhabiting (i.e., epiphyllous) bryophyte species transplanted from continuous forest into a network of 10 study sites ranging from 1, 10, and 100 to > 10,000 ha in size. All populations experienced significantly positive local growth (lambda > 1) and a nearly constant per-generational extinction probability (15%). However, experimental leaf patches in reserves of > or = 100 ha experienced nearly double (48%) the colonization probability observed in small reserves (27%), suggesting that the proximate cause of epiphyll species loss in small fragments (< or = 10 ha) is reduced colonization. Nonetheless, populations of small fragments exhibit rates of colonization above patch extinction, positive local growth, and low temporal variation, which are features that should theoretically reduce the probability of extinction. This result suggests that for habitat-tracking metapopulations subject to frequent and stochastic turnover events, including epiphylls, colonization/extinction ratios must be maintained well above unity to ensure metapopulation persistence.

Bryophyta↗

Logic for designing nature reserves for multiple species.

We examine the logic of designing nature reserves to understand better how to integrate the concepts of representativeness and persistence. Simple models of viability are used to evaluate how the expected number of species in the reserve system changes with variation in the risk of extinction among species, their rate of occurrence, and the distribution of species. The optimal size of individual reserves increased with the mean and variance of the probability of extinction among species and with the rate at which the risk of extinction declines with the cost of each reserve. In contrast, the rate of occurrence of species within reserves and their rate of accumulation with increasing reserve area had a relatively minor influence on the optimal size of reserves. Patterns of endemism were most important for the location of reserves. Including differences among species in the analysis reduced the optimal number of individual reserves (and increased the size of each) when operating under a fixed budget compared with reserve designs based on single species. A case study in the city of Melbourne, Australia, demonstrates the conservation value of small (approximately 1 ha) grassland reserves and the underrepresentation of Melbourne's volcanic plains in the region's conservation network.

Computer Simulation↗

Does red noise increase or decrease extinction risk? Single extreme events versus series of unfavorable conditions.

Recent theoretical studies have shown contrasting effects of temporal correlation of environmental fluctuations (red noise) on the risk of population extinction. It is still debated whether and under which conditions red noise increases or decreases extinction risk compared with uncorrelated (white) noise. Here, we explain the opposing effects by introducing two features of red noise time series. On the one hand, positive autocorrelation increases the probability of series of poor environmental conditions, implying increasing extinction risk. On the other hand, for a given time period, the probability of at least one extremely bad year ("catastrophe") is reduced compared with white noise, implying decreasing extinction risk. Which of these two features determines extinction risk depends on the strength of environmental fluctuations and the sensitivity of population dynamics to these fluctuations. If extreme (catastrophic) events can occur (strong noise) or sensitivity is high (overcompensatory density dependence), then temporal correlation decreases extinction risk; otherwise, it increases it. Thus, our results provide a simple explanation for the contrasting previous findings and are a crucial step toward a general understanding of the effect of noise color on extinction risk.

Animals↗

Evolutionary and ecological causes of the latitudinal diversity gradient in hylid frogs: treefrog trees unearth the roots of high tropical diversity.

Why are there more species in the tropics than in temperate regions? In recent years, this long-standing question has been addressed primarily by seeking environmental correlates of diversity. But to understand the ultimate causes of diversity patterns, we must also examine the evolutionary and biogeographic processes that directly change species numbers (i.e., speciation, extinction, and dispersal). With this perspective, we dissect the latitudinal diversity gradient in hylid frogs. We reconstruct a phylogeny for 124 hylid species, estimate divergence times and diversification rates for major clades, reconstruct biogeographic changes, and use ecological niche modeling to identify climatic variables that potentially limit dispersal. We find that hylids originated in tropical South America and spread to temperate regions only recently (leaving limited time for speciation). There is a strong relationship between the species richness of each region and when that region was colonized but not between the latitudinal positions of clades and their rates of diversification. Temperature seasonality seemingly limits dispersal of many tropical clades into temperate regions and shows significant phylogenetic conservatism. Overall, our study illustrates how two general principles (niche conservatism and the time-for-speciation effect) may help explain the latitudinal diversity gradient as well as many other diversity patterns across taxa and regions.

Animals↗

Will sympatric speciation fail due to stochastic competitive exclusion?

Sympatric speciation requires coexistence of the newly formed species. If divergence proceeds by small mutational steps, the new species utilize almost the same resources initially, and full speciation may be impeded by competitive exclusion in stochastic environments. We investigate this primarily ecological problem of sympatric speciation by studying the population dynamics of a diverging asexual population in a fluctuating environment. Correlation between species responses to environmental fluctuation is assumed to decrease with distance in trait space. Rapidly declining correlation in combination with high environmental variability may delay full speciation or even render it impossible. Stochastic extinctions impeding speciation are most likely when correlation decays faster than competition, for example, when demographic stochasticity is strong or when divergence is not accompanied by niche separation, such as in speciation driven entirely by sexual selection. Our general theoretical results show an interesting connection between short-term ecological dynamics and long-term, large-scale evolution.

Computer Simulation↗

Patch aging and the S-Allee effect: breeding system effects on the demographic response of plants to habitat fragmentation.

We used empirical and modeling approaches to examine effects of plant breeding systems on demographic responses to habitat fragmentation. Empirically, we investigated effects of local flowering plant density on pollination and of population size on mate availability in a common, self-incompatible purple coneflower, Echinacea angustifolia, growing in fragmented prairie habitat. Pollination and recruitment increased with weighted local density around individual flowering plants. This positive density dependence is an Allee effect. In addition, mean mate compatibility between pairs of plants increased with population size. Based on this empirical study, we developed an individual-based, spatially explicit demographic model that incorporates autosomal loci and an S locus. We simulated habitat fragmentation in populations identical except for their breeding system, self-incompatible (SI) or self-compatible (SC). Both populations suffered reduced reproduction in small patches because of scarcity of plants within pollination distance (potential mates) and inbreeding depression. But SI species experienced an additional, genetic contribution to the Allee effect (S-Allee effect) caused by allele loss at the S locus, which reduces mate availability, thereby decreasing reproduction. The strength of the S-Allee effect increases through time (i.e., patches age) because random genetic drift reduces S-allele richness. We investigate how patch aging influences extinction and discuss how the S-Allee effect influences communities in fragmented habitat.

Breeding↗

The paradoxical extinction: Exploring signatures of assortative mating as a possible mechanism that maintains canonical Red Wolf genetic ancestry in the American Gulf Coast canids.

Admixed genomes, particularly those with an evolutionary history of genetic exchange with an endangered or extinct species, are valued for innovative and unconventional conservation actions. Here, we show the substantial conservation value that the admixed canids of the Gulf Coast have as they retain high amounts of contemporary Red Wolf ancestry and unique genetic variation of past Red Wolf lineages (e.g. ghost ancestry). We analyzed 54,439 loci genotyped across the genome of 413 North American canids and investigated the role that assortative mating with respect to ancestry proportions played in the retention of endangered genetic variation. We report high correlations of inter-chromosomal ancestry proportions that varied with geographic location along Texas and Louisiana Gulf Coast populations, with the stronger signatures reported in the latter. We found that models of assortative mating promoted greater ancestry variance compared with random mating leading to increased efficiency of selection for Red Wolf and ghost alleles. Despite the Red Wolf being extinct in the wild, original, and ghost genomic variation persists in Gulf Coast admixed canids. We suggest two conservation strategies that value and preserve this unique and endangered genomic variation through designed breeding programs. Ultimately the incorporation of this ghost genetic variation would be valuable to boost the genetic viability of the ex situ Red Wolf breeding program, create in situ redundancy, and avoid extinction for this endemic American wolf species.

Animals↗

Morphological and molecular phylogenetic context of the angiosperms: contrasting the 'top-down' and 'bottom-up' approaches used to infer the likely characteristics of the first flowers.

Recent attempts to address the long-debated 'origin' of the angiosperms depend on a phylogenetic framework derived from a matrix of taxa versus characters; most assume that empirical rigour is proportional to the size of the matrix. Sequence-based genotypic approaches increase the number of characters (nucleotides and indels) in the matrix but are confined to the highly restricted spectrum of extant species, whereas morphology-based approaches increase the number of phylogenetically informative taxa (including fossils) at the expense of accessing only a restricted spectrum of phenotypic characters. The two approaches are currently delivering strongly contrasting hypotheses of relationship. Most molecular studies indicate that all extant gymnosperms form a natural group, suggesting surprisingly early divergence of the lineage that led to angiosperms, whereas morphology-only phylogenies indicate that a succession of (mostly extinct) gymnosperms preceded a later angiosperm origin. Causes of this conflict include: (i) the vast phenotypic and genotypic lacuna, largely reflecting pre-Cenozoic extinctions, that separates early-divergent living angiosperms from their closest relatives among the living gymnosperms; (ii) profound uncertainty regarding which (a) extant and (b) extinct angiosperms are most closely related to gymnosperms; and (iii) profound uncertainty regarding which (a) extant and (b) extinct gymnosperms are most closely related to angiosperms, and thus best serve as 'outgroups' dictating the perceived evolutionary polarity of character transitions among the early-divergent angiosperms. These factors still permit a remarkable range of contrasting, yet credible, hypotheses regarding the order of acquisition of the many phenotypic characters, reproductive and vegetative, that distinguish 'classic' angiospermy from 'classic' gymnospermy. The flower remains ill-defined and its mode (or modes) of origin remains hotly disputed; some definitions and hypotheses of evolutionary relationships preclude a role for the flower in delimiting the angiosperms. We advocate maintenance of parallel, reciprocally illuminating programmes of morphological and molecular phylogeny reconstruction, respectively supported by homology testing through additional taxa (especially fossils) and evolutionary-developmental genetic studies that explore genes potentially responsible for major phenotypic transitions.

Botany↗

Tracking ancient polyploids: a retroposon insertion reveals an extinct diploid ancestor in the polyploid origin of belladonna.

Polyploidy is a prominent process in plant evolution and adaptation, but molecular phylogenetic studies of polyploids based on DNA sequences have often been confounded by their complex gene and genome histories. We report here a retroposon insertion in the nuclear gene granule-bound starch synthase I (GBSSI or "waxy") that clearly reveals the ancient hybrid history of the medically important polyploid species belladonna (Atropa belladonna) and resolves the controversy over the taxonomic group to which it belongs, the tribe Hyoscyameae (Solanaceae). Our inferences based on the pattern of presence or absence of the retroposon insertion are corroborated by phylogenetic analyses of the GBSSI gene sequences. This case may suggest that retroposons are promising molecular markers to study polyploid evolution.

Atropa belladonna↗

Relative contribution of abundant and rare species to species-energy relationships.

A major goal of ecology is to understand spatial variation in species richness. The latter is markedly influenced by energy availability and appears to be influenced more by common species than rare ones; species-energy relationships should thus be stronger for common species. Species-energy relationships may arise because high-energy areas support more individuals, and these larger populations may buffer species from extinction. As extinction risk is a negative decelerating function of population size, this more-individuals hypothesis (MIH) predicts that rare species should respond more strongly to energy. We investigate these opposing predictions using British breeding bird data and find that, contrary to the MIH, common species contribute more to species-energy relationships than rare ones.

Animals↗

The true identity of the supposed giant fossil spider Megarachne.

Megarachne servinei from the Permo-Carboniferous Bajo de Véliz Formation of San Luis Province, Argentina (32 degrees 17'S, 65 degrees 25'E), was described as a giant mygalomorph spider ('tarantula') and, with its body length of 339mm, the largest known spider ever to have lived on Earth. Its identification as a spider was based on interpretations of the shape of the carapace, the position of the eye tubercle, the anterior protrusion of the carapace as a pair of chelicerae, and the posterior circular structure as the abdomen. X-radiography revealed possible morphology hidden in the matrix: cheliceral fangs, sternum, labium and coxae, and so a reconstruction of Megarachne as a giant spider was presented. Difficulties with the interpretation (unusual cuticular ornament, suture dividing the carapace and spade-like anterior border of the chelicera), together with non-preservation of synapomorphies of Araneae, provoked debate about its interpretation as a spider. Now, the holotype and a new specimen have become available for study. Megarachne is shown to be a bizarre eurypterid ('sea-scorpion'), similar to rare forms known from Carboniferous rocks of Scotland and South Africa, and is the most complete eurypterid so far recorded from Carboniferous strata of South America.

Animals↗