Modelling strategies for predicting the potential impacts of climate change on the natural distribution of species have often focused on the characterization of a species’ bioclimate envelope. A number of recent critiques have questioned the validity of this approach by pointing to the many factors other than climate that play an important part in determining species distributions and the dynamics of distribution changes. Such factors include biotic interactions, evolutionary change and dispersal ability. This paper reviews and evaluates criticisms of bioclimate envelope models and discusses the implications of these criticisms for the different modelling strategies employed. It is proposed that, although the complexity of the natural system presents fundamental limits to predictive modelling, the bioclimate envelope approach can provide a useful first approximation as to the potentially dramatic impact of climate change on biodiversity. However, it is stressed that the spatial scale at which these models are applied is of fundamental importance, and that model results should not be interpreted without due consideration of the limitations involved. A hierarchical modelling framework is proposed through which some of these limitations can be addressed within a broader, scale-dependent

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Predicting the impacts of climate change on the distribution of species: are bioclimate envelope models useful?

Growing interest in spatial ecology is promoting new approaches to the study of seed dispersal, one of the key processes determining the spatial structure of plant populations. Seed-dispersion patterns vary among plant species, populations and individuals, at different distances from parents, different microsites and different times. Recent field studies have made progress in elucidating the mechanisms behind these patterns and the implications of these patterns for recruitment success. Together with the development and refinement of mathematical models, this promises a deeper, more mechanistic understanding of dispersal processes and their consequences.

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Spatial patterns of seed dispersal, their determinants and consequences for recruitment.

To select candidate populations of wild species to be given priority for conservation, genetic criteria gained from the study of molecular markers may be useful. Traditionally, diversity measures such as ex- pected heterozygosity or percentage of polymorphic loci have been considered. For conservation we propose instead that priority should be given to measures of allelic richness. To standardize the results of allelic rich- ness across populations, we used the technique of rarefaction. This technique allows evaluation of the ex- pected number of different alleles among equal-sized samples drawn from several different populations. We also show how the contribution of each population to total diversity can be partitioned into two components. The first is related to the level of diversity of the population and the second to its divergence from the other populations. For conservation purposes the uniqueness of a population-in terms of its allelic composition- may be at least as important as its diversity level. These new descriptors are illustrated by means of isozyme and chloroplast DNA data obtainedfor an endangered tree species, the argan tree of Morocco (Argania spinosa (L.) Skeels). With these analyses the conservation value of the argan tree populations, especially those of two isolates present in the north of the country, can be better appreciated. The methods proposed to identify prior- ity areas for conservation of the genetic resources of the argan tree are compared to those sometimes advo- cated in the case of reserve design, where one of the goals is to maximize species richness.

Identifying Populations for Conservation on the Basis of Genetic Markers

LEARNING The objectives of BIOS 781 are to present: OBJECTIVES: 1. basic population and quantitative genetic principles, including classical genetics, chromosomal theory of inheritance, and meiotic recombination 2. an exposure to QTL mapping methods of complex quantitative traits and linkage methods to detect co-segregation with disease 3. methods for assessing marker-disease linkage disequilibrium, including case-control approaches 4. methods for genome-wide association and stratification control.

Genetic Data Analysis

The difficulty of directly measuring gene flow has lead to the common use of indirect measures extrapolated from genetic frequency data. These measures are variants of FST, a standardized measure of the genetic variance among populations, and are used to solve for Nm, the number of migrants successfully entering a population per generation. Unfortunately, the mathematical model underlying this translation makes many biologically unrealistic assumptions; real populations are very likely to violate these assumptions, such that there is often limited quantitative information to be gained about dispersal from using gene frequency data. While studies of genetic structure per se are often worthwhile, and FST is an excellent measure of the extent of this population structure, it is rare that FST can be translated into an accurate estimate of Nm.

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Indirect measures of gene flow and migration: FST≠1/(4Nm+1)

Long-distance seed dispersal influences many key aspects of the biology of plants, including spread of invasive species, metapopulation dynamics, and diversity and dynamics in plant communities. However, because long-distance seed dispersal is inherently hard to measure, there are few data sets that characterize the tails of seed dispersal curves. This paper is structured around two lines of argument. First, we argue that long-distance seed dispersal is of critical importance and, hence, that we must collect better data from the tails of seed dispersal curves. To make the case for the importance of long-distance seed dispersal, we review existing data and models of long-distance seed dispersal, focusing on situations in which seeds that travel long distances have a critical impact (colonization of islands, Holocene migrations, response to global change, metapopulation biology). Second, we argue that genetic methods provide a broadly applicable way to monitor long-distance seed dispersal; to place this argument in context, we review genetic estimates of plant migration rates. At present, several promising genetic approaches for estimating long-distance seed dispersal are under active development, including assignment methods, likelihood methods, genealogical methods, and genealogical/demographic methods. We close the paper by discussing important but as yet largely unexplored areas for future research.

Long-distance seed dispersal in plant populations.

Long-distance dispersal (LDD) includes events in which propagules arrive, but do not necessarily establish, at a site far removed from their origin. Although important in a variety of ecological contexts, the system-specific nature of LDD makes "far removed" difficult to quantify, partly, but not exclusively, because of inherent uncertainty typically involved with the highly stochastic LDD processes. We critically review the main methods employed in studies of dispersal, in order to facilitate the evaluation of their pertinence to specific aspects of LDD research. Using a novel classification framework we identify six main methodological groups: biogeographical: Eulerian and Lagrangian movement/redistributiona; short-term and long-term genetic analyses; and modeling. We briefly discuss the strengths and weaknesses of the most promising methods available for estimation of LDD. illustrating them with examples from current studies. The rarity of LDD events will continue to make collecting, analyzing, and interpreting the necessary data difficult, and a simple and comprehensive definition of LDD will remain elusive. However, considerable advances have been made in some methodological areas, such as miniaturization of tracking devices, elaboration of stable isotope and genetic analyses, and refinement of mechanistic models. Combinations of methods are increasingly used to provide improved insight on LDD from multiple angles. However, human activities substantially, increase the variety of long-distance transport avenues, making the estimation of LDD even more challenging.

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Methods for estimating long-distance dispersal

One of the major problems faced by conservation biologists is the allocation of scarce resources to an overwhelmingly large number of species in need of preservation efforts. Both demographic and genetic information have been brought to bear on this problem; however, the role of information obtained from genetic markers has largely been limited to the characterization of gene frequencies and patterns of diversity. While the genetic consequences of rarity may be a contributing factor to endangerment, it is widely recognized that demographic factors often may be more important. Because patterns of genetic marker variation are influenced by the same demographic factors of interest to the conservation biologist, it is possible to extract useful demographic information from genetic marker data. Such an approach may be productive for determining plant mating systems, inbreeding depression, effective population size, and metapopulation structure. In many cases, however, data consisting only of marker frequencies are inadequate for these purposes. Development of genealogical based analytical methods coupled with studies of DNA sequence variation within and among populations is likely to yield the most information on demographic processes from genetic marker data. Indeed, in some cases it may be the only means of obtaining information on the long-term demographic properties that may be most useful for determining the future prospects of a species of interest.

Conservation genetics: beyond the maintenance of marker diversity

The residence time of fine-root carbon in soil is one of the least understood aspects of the global carbon cycle, and fine-root dynamics are one of the least understood aspects of plant function. Most recent studies of these belowground dynamics have used one of two methodological strategies. In one approach, based on analysis of carbon isotopes, the persistence of carbon is inferred; in the other, based on direct observations of roots with cameras, the longevity of individual roots is measured. We show that the contribution of fine roots to the global carbon cycle has been overstated because observations of root lifetimes systematically overestimate the turnover of fine-root biomass. On the other hand, isotopic techniques systematically underestimate the turnover of individual roots. These differences, by virtue of the separate processes or pools measured, are irreconcilable.

https://science.sciencemag.org/content/sci/319/5862/456.full.pdf

Irreconcilable differences: Fine-root life spans and soil carbon persistence

While DNA-based markers can provide a wealth of information for the study of plant evolutionary biology, progress is limited by the lack of primers available for PCR. To overcome this limitation, we outline a protocol for developing oligonucleotide primers targeting regions of low copy-number nuclear genes. This protocol is intended to lead to universally useful primer sets. To test our approach, we designed eight primer sets and tested their abilities to amplify targets from representatives of each dicot and one monocot subclass. Five of the eight primer sets amplified targets from at least five of the seven taxa and thus exhibited broad taxonomic usefulness; the remaining primers were rather specific, however, and amplified targets from at most three taxa. In only one primer-taxon combination was a complex multiple-banded amplification produced. Overall, the protocol outlined proved quite useful at identifying broadly applicable primers targeted to low copy-number nuclear genes. Wider application of this approach should be effective at greatly increasing the amount of genetic information available for a diversity of plant nuclear genomes.

Allan E. Strand

Papers

Long-distance seed dispersal in plant populations.

Methods for estimating long-distance dispersal

Conservation genetics: beyond the maintenance of marker diversity

Irreconcilable differences: Fine-root life spans and soil carbon persistence

Nuclear DNA-based markers for plant evolutionary biology.