Height-growth response to climatic changes differs among populations of Douglas-fir: a novel analysis of historic data

Projected climate change will affect existing forests, as substantial changes are predicted to occur during their life spans. Species that have ample intraspecific genetic differentiation, such as Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), are expected to display population-specific growth responses to climate change. Using a mixed-effects modeling approach, we describe three-year height (HT) growth response to changes in climate of interior Douglas-fir populations. We incorporate climate information at the population level, yielding a model that is specific to both species and population. We use data from provenance tests from previous studies that comprised 236 populations from Idaho, Montana, and eastern Washington, USA. The most sensitive indicator of climate was the mean temperature of the coldest month. Population maximum HT and HT growth response to changes in climate were dependent on seed source climate. All populations had optimum HT growth when transferred to climates with warmer winters; those originating in sites with the warmest winters were taller across sites and had highest HT growth at transfer distances closest to zero; those from colder climates were shortest and had optimum HT growth when transferred the farthest. Although this differential response damped the height growth differences among populations, cold-climate populations still achieved their maximum growth at lower temperatures than warm-climate populations. The results highlight the relevance of understanding climate change impacts at the population level, particularly in a species with ample genetic variation among populations.     

Life style and genetic variation in teleosts: the case of pelagic (<Emphasis Type="Italic">Aphia minuta</Emphasis>) and benthic (<Emphasis Type="Italic">Gobius niger</Emphasis>) gobies (Perciformes: Gobiidae)

The pattern of genetic variability of two species of Mediterranean gobiids was compared, with reference to their different life history traits (Aphia minuta paedomorphic and pelagic; Gobius niger metamorphosed and benthic). The aim was to evaluate how different life histories can affect the genetic structure in these marine teleosts. The study was carried out on populations of both species sampled in the western Mediterranean and in the Adriatic Sea. Seven restriction endonucleases were used for the RFLP analysis of a mitochondrial DNA segment comprising the NADH dehydrogenase subunits 3, 4L and 4. The results highlighted two different patterns of genetic variation, a weak genetic structure in A. minuta and population subdivision in G. niger. These observations may be explained not only in terms of the different dispersal capabilities of these species, but also considering that A. minuta is an abbreviate iteroparous spawner while G. niger is a protracted iteroparous spawner. Because abbreviate iteroparity is a reproductive strategy selected in stable environments with high resource availability, Pliocene and Pleistocene climate oscillations may have represented factors that negatively influenced the reproductive success of A. minuta, producing demographic fluctuations and bottlenecks, as suggested by the mismatch distribution analysis. The weak genetic structure of A. minuta populations seems to be therefore due to a more recent re-colonization of the Mediterranean basin after a severe population decline, rather than to the high vagility of this pelagic goby.     

Marine extinctions and conservation

In contrast to the large number of terrestrial extinctions that have taken place over the past 12,000 years, there have apparently been very few marine extinctions. But these small losses should not be reason for complacency. During the past 50 years, government supported, commercial fishing has resulted in the collapse of about a thousand populations that once supplied most of the world’s seafood. For the collapsed species, now existing as small remnants of their former population sizes, the future is bleak. They suffer from loss of genetic diversity, inbreeding depression, and depensation. Because marine species were eliminated by historic climatic changes, continued global warming is likely to result in the extinction of small populations that already have a precarious existence. They may be considered evidence of an extinction debt that must be paid as the climate change becomes more severe. For some of the remnant species, extinction can be avoided if there is a rapid management conversion to the use of more marine protected areas (MPAs) and extensive ocean zoning where fishing is prohibited.     

Population structure of the common sole (<Emphasis Type="Italic">Solea solea</Emphasis>) in the Northeastern Atlantic and the Mediterranean Sea: revisiting the divide with EPIC markers

Spatial and temporal population genetic structures of the common sole, Solea solea, were studied in Northeastern Atlantic and Mediterranean Sea populations, using three polymorphic exon-primed intron-crossing (EPIC) markers. Results demonstrated significant multilocus differentiation among Eastern Mediterranean and a group composed by Western Mediterranean and Atlantic populations (θ = 0.150, P < 0.001), but also suggested unrecorded genetic differentiation of the Adriatic Sea population. No pattern of isolation-by-distance was recorded across the range covered by sampling, from the Kattegat to the Aegean Sea. Conversely to genetically structured Mediterranean populations, Atlantic populations ranging from Denmark to Portugal could be considered as representative of the same panmictic unit (θ = 0.009, not significant). Results further demonstrated stability of multilocus genetic structure among temporarily replicated cohort samples [0+, 1+, subadults] from several coastal and estuarine locations from Bay of Biscay, excepted for the amylase locus Am2B3-2 at one location (Pertuis d’Antioche). Despite coherence of such observed patterns of multilocus differentiation with previous allozymic surveys in sole, and with patterns generally obtained for other marine fish species, single-locus results from EPICs indicated divergent coalescence schemes supporting a complex response to ecology and history of sole’s populations. Results stress the use of nuclear genes such as EPIC markers to investigate population structure, but also historical, demographic, and possibly selective processes in marine fishes.     

Genetic structure and allozyme variation of sea bass (Dicentrarchus labrax and D. punctatus) in the Mediterranean Sea

This paper reports data on 28 allozyme loci in wild and artificially reared sea bass (Dicentrarchus labrax) samples, originating from either coastal lagoon or marine sites in the Mediterranean Sea. F _ST analysis (θ estimator) indicated strong genetic structuring among populations; around 34% of the overall genetic variation is due to interpopulation variation. Pairwise θ estimates showed that, on average, the degree of genetic structuring was much higher between marine populations than between samples from lagoons. Six polymorphic loci showed differences in allele frequencies between marine and lagoon samples. Multivariate analyses of individual allozymic profiles and of allele frequencies suggested that different arrays of genotypes prevail in lagoons compared to marine samples, particularly at those loci that, on the basis of previous acclimation experiments, had been implicated in adaptation to freshwater. On the other hand, variation at “neutral” allozyme loci reflects to a greater extent the geographic location of populations. Allozyme differentiation was also studied in a D. labrax population from the Portuguese coast. Average genetic distance between this population and the Mediterranean populations was quite high (Nei's D = 0.236) and calls into question the taxonomic status of the Portuguese population. Finally, genetic relationships between D. labrax and D. punctatus were evaluated. Average Nei's D was 0.648, revealing high genetic differentiation between the two species, even for two sympatric populations of these species in Egypt; thus gene flow was not indicated between species. Received: 24 October 1996 / Accepted: 27 November 1996     

Fire Management,Managed Relocation,and Land Conservation Options for Long‐Lived Obligate Seeding Plants under Global Changes in Climate,Urbanization, and Fire Regime

Most species face multiple anthropogenic disruptions. Few studies have quantified the cumulative influence of multiple threats on species of conservation concern, and far fewer have quantified the potential relative value of multiple conservation interventions in light of these threats. We linked spatial distribution and population viability models to explore conservation interventions under projected climate change, urbanization, and changes in fire regime on a long‐lived obligate seeding plant species sensitive to high fire frequencies, a dominant plant functional type in many fire‐prone ecosystems, including the biodiversity hotspots of Mediterranean‐type ecosystems. First, we investigated the relative risk of population decline for plant populations in landscapes with and without land protection under an existing habitat conservation plan. Second, we modeled the effectiveness of relocating both seedlings and seeds from a large patch with predicted declines in habitat area to 2 unoccupied recipient patches with increasing habitat area under 2 projected climate change scenarios. Finally, we modeled 8 fire return intervals (FRIs) approximating the outcomes of different management strategies that effectively control fire frequency. Invariably, long‐lived obligate seeding populations remained viable only when FRIs were maintained at or above a minimum level. Land conservation and seedling relocation efforts lessened the impact of climate change and land‐use change on obligate seeding populations to differing degrees depending on the climate change scenario, but neither of these efforts was as generally effective as frequent translocation of seeds. While none of the modeled strategies fully compensated for the effects of land‐use and climate change, an integrative approach managing multiple threats may diminish population declines for species in complex landscapes. Conservation plans designed to mitigate the impacts of a single threat are likely to fail if additional threats are ignored. Manejo de Incendios, Reubicación Administrada y Opciones de Conservación de Suelo para Plantas de Vida Larga con Sembrado Obligado bajo los Cambios Globales en el Clima, la Urbanización y el Régimen de Incendios     

Incorporating differences between genetic diversity of trees and herbaceous plants in conservation strategies

Reviews that summarize the genetic diversity of plant species in relation to their life history and ecological traits show that forest trees have more genetic diversity at population and species levels than annuals or herbaceous perennials. In addition, among-population genetic differentiation is significantly lower in trees than in most herbaceous perennials and annuals. Possible reasons for these differences between trees and herbaceous perennials and annuals have not been discussed critically. Several traits, such as high rates of outcrossing, long-distance pollen and seed dispersal, large effective population sizes (N_e), arborescent stature, low population density, longevity, overlapping generations, and occurrence in late successional communities, may make trees less sensitive to genetic bottlenecks and more resistant to habitat fragmentation or climate change. We recommend that guidelines for genetic conservation strategies be designed differently for tree species versus other types of plant species. Because most tree species fit an LH scenario (low [L] genetic differentiation and high [H] genetic diversity), tree seeds could be sourced from a few populations distributed across the species’ range. For the in situ conservation of trees, translocation is a viable option to increase N_e. In contrast, rare herbaceous understory species are frequently HL (high differentiation and low diversity) species. Under the HL scenario, seeds should be taken from many populations with high genetic diversity. In situ conservation efforts for herbaceous plants should focus on protecting habitats because the typically small populations of these species are vulnerable to the loss of genetic diversity. The robust allozyme genetic diversity databases could be used to develop conservation strategies for species lacking genetic information. As a case study of reforestation with several tree species in denuded areas on the Korean Peninsula, we recommend the selection of local genotypes as suitable sources to prevent adverse effects and to insure the successful restoration in the long term.     

Applying assessments of adaptive capacity to inform natural-resource management in a changing climate

Adaptive capacity (AC)—the ability of a species to cope with or accommodate climate change—is a critical determinant of species vulnerability. Using information on species’ AC in conservation planning is key to ensuring successful outcomes. We identified connections between a list of species’ attributes (e.g., traits, population metrics, and behaviors) that were recently proposed for assessing species’ AC and management actions that may enhance AC for species at risk of extinction. Management actions were identified based on evidence from the literature, a review of actions used in other climate adaptation guidance, and our collective experience in diverse fields of global-change ecology and climate adaptation. Selected management actions support the general AC pathways of persist in place or shift in space, in response to contemporary climate change. Some actions, such as genetic manipulations, can be used to directly alter the ability of species to cope with climate change, whereas other actions can indirectly enhance AC by addressing ecological or anthropogenic constraints on the expression of a species’ innate abilities to adapt. Ours is the first synthesis of potential management actions directly linked to AC. Focusing on AC attributes helps improve understanding of how and why aspects of climate are affecting organisms, as well as the mechanisms by which management interventions affect a species’ AC and climate change vulnerability. Adaptive-capacity-informed climate adaptation is needed to build connections among the causes of vulnerability, AC, and proposed management actions that can facilitate AC and reduce vulnerability in support of evolving conservation paradigms.     

Adaptive introgression as a resource for management and genetic conservation in a changing climate

Current rates of climate change require organisms to respond through migration, phenotypic plasticity, or genetic changes via adaptation. We focused on questions regarding species’ and populations’ ability to respond to climate change through adaptation. Specifically, the role adaptive introgression, movement of genetic material from the genome of 1 species into the genome of another through repeated interbreeding, may play in increasing species’ ability to respond to a changing climate. Such interspecific gene flow may mediate extinction risk or consequences of limited adaptive potential that result from standing genetic variation and mutation alone, enabling a quicker demographic recovery in response to changing environments. Despite the near dismissal of the potential benefits of hybridization by conservation practitioners, we examined a number of case studies across different taxa that suggest gene flow between sympatric or parapatric sister species or within species that exhibit strong ecotypic differentiation may represent an underutilized management option to conserve evolutionary potential in a changing environment. This will be particularly true where advanced‐generation hybrids exhibit adaptive traits outside the parental phenotypic range, a phenomenon known as transgressive segregation. The ideas presented in this essay are meant to provoke discussion regarding how we maintain evolutionary potential, the conservation value of natural hybrid zones, and consideration of their important role in adaptation to climate.     

Evidence of Local Adaptation in the Demographic Response of American Ginseng to Interannual Temperature Variation

Abstract: Bioclimatic envelope models of species’ responses to climate change are used to predict how species will respond to increasing temperatures. These models are frequently based on the assumption that the northern and southern boundaries of a species’ range define its thermal niche. However, this assumption may be violated if populations are adapted to local temperature regimes and have evolved population‐specific thermal optima. Considering the prevalence of local adaptation, the assumption of a species‐wide thermal optimum may be violated for many species. We used spatially and temporally extensive demographic data for American ginseng (Panax quinquefolius L.) to examine range‐wide variation in response of population growth rate (λ) to climatic factors. Our results suggest adaptation to local temperature, but not precipitation. For each population, λ was maximized when annual temperatures were similar to site‐specific, long‐term mean temperatures. Populations from disparate climatic zones responded differently to temperature variation, and there was a linear relation between population‐level thermal optima and the 30‐year mean temperature at each site. For species that are locally adapted to temperature, bioclimatic envelope models may underestimate the extent to which increasing temperatures will decrease population growth rate. Because any directional change from long‐term mean temperatures will decrease population growth rates, all populations throughout a species’ range will be adversely affected by temperature increase, not just populations at southern and low‐elevation boundaries. Additionally, when a species’ local thermal niche is narrower than its range‐wide thermal niche, a smaller temperature increase than would be predicted by bioclimatic envelope approaches may be sufficient to decrease population growth.     

Two markers and one history: phylogeography of the edible common sea urchin Paracentrotus lividus in the Lusitanian region

Benthic marine invertebrates with long-lived larvae are believed to have dispersal capabilities that contribute to maintaining genetic uniformity among populations over large geographical scales. However, both hydrological and biological factors may limit the actual dispersal of such larvae. We studied the population genetic structure of the edible common sea urchin Paracentrotus lividus (Lamarck, 1816), to explore its dispersal patterns in the Atlanto-Mediterranean region and, more specifically, to ascertain the role of the Strait of Gibraltar in shaping the genetic structure of this species. For this purpose, we analysed 158 individuals for the mitochondrial 16S rRNA gene and 151 of these for the nuclear single-copy intron adenine nucleotide transporter (ANT) from 16 localities from the Atlantic and Mediterranean basins, spanning over 4,000 km. Mitochondrial 16S rRNA shows higher genetic diversity in the Mediterranean than in the Atlantic and reveals a sharp break between the populations of both basins, probably as a consequence of the barrier imposed by the Almería–Orán hydrological front, situated east of the Strait of Gibraltar. Both markers suggest that a recent population expansion has taken place in both basins, most probably following the Messinian salinity crisis. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Implications of Macroalgal Isolation by Distance for Networks of Marine Protected Areas

The global extent of macroalgal forests is declining, greatly affecting marine biodiversity at broad scales through the effects macroalgae have on ecosystem processes, habitat provision, and food web support. Networks of marine protected areas comprise one potential tool that may safeguard gene flow among macroalgal populations in the face of increasing population fragmentation caused by pollution, habitat modification, climate change, algal harvesting, trophic cascades, and other anthropogenic stressors. Optimal design of protected area networks requires knowledge of effective dispersal distances for a range of macroalgae. We conducted a global meta‐analysis based on data in the published literature to determine the generality of relation between genetic differentiation and geographic distance among macroalgal populations. We also examined whether spatial genetic variation differed significantly with respect to higher taxon, life history, and habitat characteristics. We found clear evidence of population isolation by distance across a multitude of macroalgal species. Genetic and geographic distance were positively correlated across 49 studies; a modal distance of 50–100 km maintained F_ST < 0.2. This relation was consistent for all algal divisions, life cycles, habitats, and molecular marker classes investigated. Incorporating knowledge of the spatial scales of gene flow into the design of marine protected area networks will help moderate anthropogenic increases in population isolation and inbreeding and contribute to the resilience of macroalgal forests. Implicaciones del Aislamiento por Distancia de Macroalgas para Redes de Áreas Marinas Protegidas     

Population differentiation in megrim (<Emphasis Type="Italic">Lepidorhombus whiffiagonis</Emphasis>) and four spotted megrim (<Emphasis Type="Italic">Lepidorhombus boscii</Emphasis>) across Atlantic and Mediterranean waters and implications for wild stock management

Megrim, Lepidorhombus whiffiagonis, and four spot megrim, Lepidorhombus boscii, are two marine fish species of high commercial interest. Despite their quite heavy exploitation little is known on the genetic structure of their populations. The present work aimed at characterizing the first seven microsatellites markers available for the two megrim species. These new markers were in a second step employed to describe the population structure of the two species among their almost entire habitat range (Atlantic and Mediterranean samples). Our study confirmed the existence of a strong genetic difference between Atlantic and Mediterranean megrim species already described in the literature for L. whiffiagonis on the basis of variations at ribosomal genes. Additionally our analysis gave the first evidences of a strong genetic differentiation among Atlantic populations in both megrim species (within Atlantic global F_ST in L. whiffiagonis and L. boscii were respectively 0.158 and 0.145). When describing megrim population structure, the comparison between allele-frequency-based tests (F_ST comparisons) and genotype-based inferences (Bayesian approach) gave evidences of a hierarchical structure of the populations. In conclusion, our work enlighten the existence of two different stocks within the Atlantic Ocean and one in the Mediterranean Sea that will clearly need to be managed separately. As the present results do not fully support the current megrim stock boundaries they will surely help to rethink megrim management policies in the future.     

Interacting Effects of Phenotypic Plasticity and Evolution on Population Persistence in a Changing Climate

Abstract: Climate change affects individual organisms by altering development, physiology, behavior, and fitness, and populations by altering genetic and phenotypic composition, vital rates, and dynamics. We sought to clarify how selection, phenotypic plasticity, and demography are linked in the context of climate change. On the basis of theory and results of recent empirical studies of plants and animals, we believe the ecological and evolutionary issues relevant to population persistence as climate changes are the rate, type, magnitude, and spatial pattern of climate‐induced abiotic and biotic change; generation time and life history of the organism; extent and type of phenotypic plasticity; amount and distribution of adaptive genetic variation across space and time; dispersal potential; and size and connectivity of subpopulations. An understanding of limits to plasticity and evolutionary potential across traits, populations, and species and feedbacks between adaptive and demographic responses is lacking. Integrated knowledge of coupled ecological and evolutionary mechanisms will increase understanding of the resilience and probabilities of persistence of populations and species.     

Phylogeography of the Antarctic planktotrophic brittle star Ophionotus victoriae reveals genetic structure inconsistent with early life history

In the marine environment, connectivity is influenced by physical oceanography as well as life history and behavioral traits, which in combination with historical events such as geology, physical oceanography, and climate, determine population structure. The Antarctic brittle star Ophionotus victoriae develops via a feeding planktonic larval stage, and therefore has potential for long-distance dispersal throughout its Antarctic/subantarctic range. To evaluate this hypothesis, phylogeography of this ecologically dominant species was elucidated by sequence analysis of two mtDNA genes from individuals collected throughout the Antarctic Peninsula and from two subantarctic islands. Counter to expectations of genetic homogeneity, mtDNA data revealed substantial levels of genetic differentiation as well as diversity. Although there were some genetically homogeneous populations, such as those throughout Bransfield Strait, we found O. victoriae to have significant population structure throughout much of the Antarctic Peninsula, with evidence of potential cryptic speciation between the western and eastern Antarctic Peninsula. Furthermore, Antarctic Peninsula populations were genetically distinct from subantarctic island populations. The low levels of connectivity implied for O. victoriae contrast with those found for many other Antarctic benthic taxa, and suggest a complex interplay between oceanography, recent climate history, and larval ecology.     

Climate Change,Marine Environments,and the U.S. Endangered Species Act

Climate change is expected to be a top driver of global biodiversity loss in the 21st century. It poses new challenges to conserving and managing imperiled species, particularly in marine and estuarine ecosystems. The use of climate‐related science in statutorily driven species management, such as under the U.S. Endangered Species Act (ESA), is in its early stages. This article provides an overview of ESA processes, with emphasis on the mandate to the National Marine Fisheries Service (NMFS) to manage listed marine, estuarine, and anadromous species. Although the ESA is specific to the United States, its requirements are broadly relevant to conservation planning. Under the ESA, species, subspecies, and “distinct population segments” may be listed as either endangered or threatened, and taking of most listed species (harassing, harming, pursuing, wounding, killing, or capturing) is prohibited unless specifically authorized via a case‐by‐case permit process. Government agencies, in addition to avoiding take, must ensure that actions they fund, authorize, or conduct are not likely to jeopardize a listed species’ continued existence or adversely affect designated critical habitat. Decisions for which climate change is likely to be a key factor include: determining whether a species should be listed under the ESA, designating critical habitat areas, developing species recovery plans, and predicting whether effects of proposed human activities will be compatible with ESA‐listed species’ survival and recovery. Scientific analyses that underlie these critical conservation decisions include risk assessment, long‐term recovery planning, defining environmental baselines, predicting distribution, and defining appropriate temporal and spatial scales. Although specific guidance is still evolving, it is clear that the unprecedented changes in global ecosystems brought about by climate change necessitate new information and approaches to conservation of imperiled species. El Cambio Climático, los Ecosistemas Marinos y el Acta Estadunidense de Especies en Peligro     

Mathematical modelling for conservation and management of gorgonians corals: youngs and olds,could they coexist?

Gorgonian corals are long-lived, slow-growing marine species dominating Mediterranean rocky bottoms. Endowed with complex morphologies they give a structure to the whole community, moreover, being efficient suspension feeders, they play a key role in plankton-benthos energy flow and CO₂${\text{CO}}_2$ storage. Thus, the structure and the development of benthic, hard bottom communities are linked to gorgonian survival. The red coral Corallium rubrum (L. 1758) is a precious gorgonian endemic to the Mediterranean Sea. Harvested and traded world-wide since ancient times red coral is a clear example of overexploited marine resource. This species is structured into self-seeding, genetically differentiated populations, some of which, living in the shallower part of the species bathymetric distribution, was recently affected by anomalous mortality events linked to global climate change. The co-occurrence of overharvesting and mass mortality could dramatically affect such populations. Demographic population models, widely applied by conservation biologists to check population viability and to project population trends over time are fundamental to foster survival of such populations matching harvesting to population growth rates. Therefore we set out a dynamic model of a genetically differentiated red coral population living in shallow waters. This population is characterised by small/young, crowded colonies and high recruitment rate. On the basis of the size–age structure determined for this population, a static life-history table, in which survival and reproduction coefficients of the different size–age classes were reported, has been set out. Demographic data were included in a non-linear, discrete, age-structured dynamic model, based on a Leslie-Lewis transition matrix. Our field data indicate that the recruits-to-larvae ratio is actually density-dependent. Such dependence, positive for low and negative for high density values, was included into the model and the effect of colonies of different size–age classes on recruits-to-larvae ratio was considered to be proportional to the number of polyps they have. We applied such model to simulate the trends of the studied population under different increases of survival and life-span. As some populations of gorgonians actually show the dominance of sparse, big/old colonies and low recruitment rate, while others are characterised by crowded, small/young colonies and high recruitment rate, we simulated the shift from the former to the latter structure increasing survival and life-span. Our results suggest that a dramatic mortality increase of bigger–older colonies (due, in the case of red coral to overfishing) could have determined the population structure we found.     

The immediate costs and long-term benefits of assisted gene flow in large populations

With the genetic health of many plant and animal populations deteriorating due to climate change outpacing adaptation, interventions, such as assisted gene flow (AGF), may provide genetic variation necessary for populations to adapt to climate change. We ran genetic simulations to mimic different AGF scenarios in large populations and measured their outcomes on population-level fitness to determine circumstances in which it is worthwhile to perform AGF. In the absence of inbreeding depression, AGF was beneficial within a few generations only when introduced genotypes had much higher fitness than local individuals and traits affecting fitness were controlled by a few genes of large effect. AGF was harmful over short periods (e.g., first ∼10–20 generations) if there was strong outbreeding depression or introduced deleterious genetic variation. When the adaptive trait was controlled by many loci of small effect, the benefits of AGF took over 10 generations to realize—potentially too long for most climate-related management scenarios. The genomic integrity of the recipient population typically remained intact following AGF; the amount of genetic material from the donor population usually constituted no more of the recipient population's genome than the fraction of the population introduced. Significant genomic turnover (e.g., >50% replacement) only occurred when the selective advantage of the adaptive trait and translocation fraction were extremely high. Our results will be useful when adaptive management is used to maintain the genetic health and productivity of large populations under climate change.