Habitat fragmentation effects on fitness of plant populations – a review

doi:10.1016/j.jnc.2003.07.002

Journal for Nature Conservation

Volume 12, Issue 1, 2 July 2004, Pages 53-72

https://doi.org/10.1016/j.jnc.2003.07.002 Get rights and content

Abstract

Habitat fragmentation threatens the survival of many species and local populations. Habitat fragmentation has two major consequences: populations become more isolated and are reduced in size. Small compared with large populations have increased extinction risks because of different types stochasticity (e.g. genetic drift) and inbreeding, which can negatively affect the fitness of individuals or populations. Habitat fragmentation may also change the abiotic conditions of the surrounding landscape, which influences biotic interactions. This review gives an introduction to the theory of the effects of habitat fragmentation on mean fitness of plant populations. It intends to help bridge the gap between conservation biologists and conservation practitioners. The paper shortly introduces basic concepts of population biology, demography and genetics and cites relevant and new literature. Special attention is given to more common plant species, which have attracted far less conservation attention than rare species.

Section snippets

Habitat fragmentation

Human activities have changed between one-third and one-half of the earth's land surface and are leading to substantial and growing modification of the earth's biological resources (Vitousek, Mooney, Lubchenco, & Melillo, 1997). Among others, habitat destruction and habitat fragmentation are ongoing major anthropogenic impacts on landscapes, which can strongly affect ecosystems, populations and species (Young, Boyle, & Brown, 1996; Young & Clarke, 2000). Plants and animals may live in naturally

What is a metapopulation?

Scattered populations of a species in a landscape, which are in contact via migration and which are characterised by local population extinction and by colonisation of unoccupied sites constitute a metapopulation (Levins, 1969; Hanski & Gilpin (1991), Hanski & Gilpin (1997)). Especially in the light of habitat fragmentation, the metapopulation concept has received increasing interest, and it has been widely discussed and expanded (e.g. Gotelli, 1991; Hanski & Gilpin (1991), Hanski & Gilpin

Natural catastrophes and environmental stochasticity

Habitat fragmentation creates small and/or isolated populations. Such populations have increased extinction risks by pure chance, compared with larger populations. Four types of stochasticity affecting populations are recognised: natural catastrophes, environmental stochasticity, demographic stochasticity and genetic drift (Shaffer, 1987). Natural catastrophes such as floods, fires or droughts can, of course, also drive large populations or even species to extinction in short time. However,

Genetic mechanisms and inbreeding depression

The term ‘inbreeding’ is used to describe various related phenomena; all of them refer to matings among relatives, which increases homozygosity, and all of them are relative measures (see Keller & Waller, 2002). (a) ‘Pedigree inbreeding’ means that an individual is inbred if parents share ancestors; or in other words that two genes in different individuals are derived from the same gene in a common ancestor. (b) Inbreeding can be a result of non-random mating, meaning that the parents of an

Mutations and purging

Mutations occur randomly in any individual. Often mutations have no consequences, because they occur in unimportant regions in the genome (e.g. in spacer regions, which are not translated into proteins and are not expressed). If mutations do have consequences, they are mostly negative and can be mildly deleterious to lethal. But mutations are also the source of genetic variation, and theoretically balance the loss of genetic variability through genetic drift (Lande, 1995). Moreover, mutations

Number of migrants

The negative effects of small population size on genetic mechanisms described above (i.e. the effects of genetic drift on neutral and non-neutral genetic diversity) can be alleviated if the populations are not completely isolated and receive additional ‘genetic material’ from other populations. This is called gene flow (Levin & Kerster, 1974; Slatkin, 1985; Levin, 1988; Ellstrand (1992a), Ellstrand (1992b); Ellstrand & Elam, 1993). The importance and magnitude of gene flow has been widely

Plant–pollinator interactions

Apart from the genetic effects described above, different reactions of pollinators in small compared with large populations may also reduce population viability. In small plant patches and in patches with reduced plant densities, pollinator diversity is often reduced (Rathcke & Jules, 1993). Often, pollinators are only attracted to large patches of the one same species or to large patches with many different flowers growing on them (facilitation effect; Sih & Baltus, 1987; Olesen & Jain, 1994).

Rare plants

Through habitat fragmentation, plants (and animals) can be forced to live in small populations. This may make them rarer than they were before and can eventually lead to local population or even species extinction (Young et al., 1996; Young & Clarke, 2000). On the other hand, many plant species have evolved over long time ranges in naturally small, isolated populations and are well adapted to surviving under these special conditions. Several types of rarity are acknowledged in plant ecology:

Altered landscape structure and edge effects

Habitat fragmentation may not only have a direct impact on populations, by dividing and isolating them, but it can also change abiotic conditions of the surrounding landscape and of the habitat itself (Saunders et al., 1991). This will inevitably influence biotic interactions in the habitat remnant. For instance, if a certain habitat was formerly surrounded by woodland, wind flux across the landscape may be altered after destruction of the woodland. Such an effect was found in the maple Acer

Minimum viable population sizes

For practicing conservation managers in particular, an important question arising from the above-described negative effects of small population size is how large a viable population would have to be. Much scientific effort has been placed into determining minimum viable population sizes (MVP). Widely accepted mathematical models showed that populations with less than 100–500 reproductive individuals will not be able to survive in medium time ranges (say around 100 years), because of chance

Artificial gene flow and outbreeding depression

Since gene flow can alleviate the problems associated with isolation of small populations, it has been suggested to use artificial gene flow as a conservation measure. For instance, seeds from one large and vigorous population could be brought to a small and endangered population, hereby increasing its genetic diversity. However, gene flow is not necessarily only beneficial. Gene flow can also reduce fitness, because sessile plants with restricted dispersal capacities may be highly adapted to

Summary and conclusions

Habitat fragmentation threatens the survival of many species and can cause the extinction of local populations (Ehrlich & Wilson, 1991; Young et al., 1996; Young & Clarke, 2000). Habitat fragmentation has two major consequences: populations become more isolated and are reduced in size. Small compared with large populations have increased extinction risks because of natural catastrophes and environmental, demographic or genetic stochasticity (drift). Genetic drift reduces the genetic variability

Acknowledgements

The author thanks Elena Conti, Matthias Diemer, Markus Fischer, Bernhard Schmid, Jakob Schneller and especially Isabelle Olivieri for critically reading earlier versions of the manuscript, for helpful comments and for their encouragement.

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Citation Excerpt :
Such fragmentation-dependent effects on abiotic and biotic environmental characteristics may also pose selection on dispersal-related traits, resulting in additional indirect paths through which fragmentation affects dispersal. For example, fragmentation is often associated with reduction in plant population sizes and with reduced accessibility for pollinators (Lázaro et al., 2020; Lienert, 2004); eventually leading to reduced genetic diversity and increased inbreeding load (Legrand et al., 2017). These changes can subsequently elevate levels of inbreeding depression in the population, a process which in many cases is associated with reduced seed mass.
Habitat fragmentation entails major effects on many ecological processes. Theory offers two contradicting hypotheses for the expected effects of fragmentation on seed dispersal strategy. On the one hand, fragmentation may select for increased dispersal by purging poor dispersers that are incapable of moving between patches (spatial sorting). On the other hand, fragmentation may select for reduced dispersal due to the increased cost of leaving a colonized patch; especially whenever the surrounding matrix is hostile. Furthermore, fragmentation can have additional indirect effects on dispersal through its' effects on abiotic and biotic environmental characteristics, but these pathways are rarely accounted for. Empirical evidence testing these hypotheses in plants comes mainly from distinctly fragmented landscapes at either very large scales or very small scales, whereas studies at intermediate scales, such as those that characterize many agricultural landscapes are lacking. Here, we quantified direct and indirect effects of fragmentation-related characteristics, such as patch size and isolation level, on multiple dispersal-related traits of a heterocarpic annual plant in a fragmented agricultural landscape. Using a structural equation modelling approach, we examined whether fragmentation had positive or negative effects on seed dispersal potential and seed dispersal-related traits. Overall, we found more support for the spatial sorting hypothesis. However, the patterns were not simple, nor consistent across all traits and paths. The positive association between fragmentation and dispersal was mainly via a negative effect of connectivity on seed dispersal potential, but the effect of patch area on dispersal traits was less clear. We identified multiple direct and indirect effects of fragmentation-related patch characteristics on dispersal-related traits. Furthermore, the relative importance of direct and indirect effects was qualitatively different between overall effects. Our findings show that different characteristics of fragmented habitats may differentially affect different dispersal-related traits, and that these effects may take place via multiple paths, often leading to a complex relationship between fragmentation and dispersal.

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Habitat fragmentation effects on fitness of plant populations – a review

Abstract

Section snippets

Habitat fragmentation

What is a metapopulation?

Natural catastrophes and environmental stochasticity

Genetic mechanisms and inbreeding depression

Mutations and purging

Number of migrants

Plant–pollinator interactions

Rare plants

Altered landscape structure and edge effects

Minimum viable population sizes

Artificial gene flow and outbreeding depression

Summary and conclusions

Acknowledgements

Trends in Ecology and Evolution

Trends in Ecology and Evolution

Journal of Theoretical Biology

Biological Conservation

Biological Journal of the Linnean Society

Biological Journal of the Linnean Society

Trends in Ecology and Evolution

Trends in Ecology and Evolution

Biological Conservation

Basic and Applied Ecology

Population structure and morph-specific differences in tristylous Lythrum salicaria

Evolution

Forest fragmentation, pollination, and plant reproduction in a chaco dry forest, Argentina

Ecology

Principles of animal ecology

Isolation, gene flow, and genetic differentiation among populations

Ecological genetics of metapopulationsThe Silene-Ustilago plant–pathogen system

Effects of host plant patch size on herbivore densityPatterns

Ecology

Evolution and maintenance of stigma-height dimorphism in Narcissus. I. Floral variation and style-morph ratios

Heredity

Heterostylous genetic polymorphismsmodel systems for evolutionary analysis

Effects of a change in the level of inbreeding on the genetic load

Nature

Genetic and evolutionary consequences of small population size in plantsimplications for conservation

The dissolution of a complex genetic polymorphismThe evolution of self-fertilization in tristylous Eichhornia paniculata (pontederiaceae)

Evolution

Inbreeding depression due to mildly deleterious mutations in finite populationsSize does matter

Genetical Research

EcologyIndividuals, populations and communities

Genetic diversity and the survival of populations

Plant Biology

Population viability analysis

Annual Review of Ecology and Systematics

Long distance seed dispersal by windMeasuring and modelling the tail of the curve

Oecologia

Temporal and spatial changes in a metapopulation of the rust pathogen Triphragmium ulmariae and its host, Filipendula ulmaria

Journal of Ecology

Coevolution at multiple spatial scalesLinum marginale–Melampsoralini—from the individual to the species

Evolutionary Ecology

Do plant populations purge their genetic load? Effects of population size and mating history on inbreeding depression

Annual Review of Ecology and Systematics

Four-year growth and reproduction of Cyclopogon cranichoides (Orchidaceae) in South Florida

American Journal of Botany

Pollen and gene dispersalThe influences of competition for pollination

Evolution

Infection by pathogens and population age of host plants

Journal of Ecology

The spatial distribution of plant populations, disease dynamics and evolution of resistance

Oikos

Inbreeding depression and its evolutionary consequences

Annual Review of Ecology and Systematics

The maintenance of nucleocytoplasmic polymorphism in a metapopulationThe case of gynodioecy

The American Naturalist

The evolution of generation time in metapopulations of monocarpic perennial plantsSome theoretical considerations and the example of the rare thistle Carlina vulgaris

Evolutionary Ecology

Inbreeding and outbreeding depression in fragmented populations

Stochastic loss of style morphs from populations of tristylous Lythrum salicaria and Decodon verticillatus (Lythraceae)

Evolution

Biodiversity studiesScience and policy

Science

Vegetation Mitteleuropas mit den Alpen in ökologischer dynamischer und historischer Sicht

Gene flow by pollenImplications for plant conservation genetics

Oikos