Impacts of Habitat Loss and Fragmentation On Terrestrial Biodiversity

Impacts of Habitat Loss and Fragmentation on Terrestrial Biodiversity
Jordan E Rogan, Texas A&M University, College Station, TX, United States
Thomas E Lacher Jr, Texas A&M University, College Station, TX, United States; Global Wildlife Conservation, Austin, TX, United States
© 2018 Elsevier Inc. All rights reserved.
Introduction: Impacts of Habitat Loss and Fragmentation on Terrestrial Biodiversity 1

The Theory of Island Biogeography 2
SLOSS: Single Large or Several Small? 2
Defining and Distinguishing Habitat Loss From Habitat Fragmentation 2
Threshold Hypotheses 3
Main Drivers of Habitat Loss and Fragmentation 4
Agriculture and Food Production 4
Logging, Deforestation and Degradation 4
Urban Expansion 4
Impacts of Habitat Loss and Fragmentation on Terrestrial Biodiversity 4
Habitat Loss 4
Habitat Fragmentation 4
Influence of Habitat Fragmentation 5
Habitat area 5
Fragment shape 7
Matrix influence 7
Edge effects 8
Isolation 9
Confounding Factors in Drawing Generalizations on the Influence of Habitat Loss and Fragmentation 10
Spatial Scale of Analysis 10
Nonlinearity in Species Response 10
Temperate Versus Tropical Systems 11
Synergistic Interactions Between Fragmentation and Other Threatening Processes 11
Time Lags 12
Lessons From Large-Scale, Long-Term Fragmentation Experiments 12
Conclusions and Future Directions 14
References 15
Introduction: Impacts of Habitat Loss and Fragmentation on Terrestrial Biodiversity
Habitat loss and fragmentation are widely recognized as the key drivers of global biodiversity loss occurring today (Rands et al.,
2010). Biodiversity can be described as the variety of genes, species, and ecosystems that constitute life on Earth (Rands et al., 2010).
Thereby it is a concept that is used to guide conservation planning, particularly when evaluating the influence of habitat disturbance
on species populations (Primack, 2014). The Millennium Ecosystem Assessment (MEA, 2005a,b) determined that over half of the
world’s biomes, such as temperate forests, tropical and subtropical dry broadleaf forests, and the Mediterranean, had been
converted by the year 1990. New satellite imagery has begun to reveal the extent to which people are restructuring ecosystems
across the globe. Nearly 43% of the terrestrial world surface has been converted from its natural state for anthropogenic purposes
(Barnosky et al., 2012). Prevalent throughout temperate systems between the 1700 and 1900s, deforestation has only increased its
pervasiveness into tropical regions in the last 100 years, leading to a loss of over one third of global forest cover (Haddad et al.,
2015). Over 50% of tropical forests have been degraded globally (Chazdon et al., 2009), and only 2–3% of original forests remain
in natural or natural-like state in Western Europe (WWF Report, 2001) (Fig. 1).
Global assessments such as these have shown that these landscape processes are resulting in declining population sizes and
increasing extinction risk of many terrestrial species (Tittensor et al., 2014). Studies investigating the influence of habitat loss and
fragmentation on biodiversity and species extinction risk have been a major focus of biodiversity conservation research (e.g.,
Andren, 1994; Fahrig, 1997; Flather and Bevers, 2002; With and King, 1999, etc.) and are therefore prevalent topics in current
scientific literature. Habitat fragmentation typically results from overall habitat loss and refers to the decline of continuous tracts of
habitat to smaller, more isolated remnant patches (Collinge, 2009). While there is continued debate regarding how to differentiate
habitat loss from fragmentation and measure their relative importance (Didham, 2010; Fahrig, 2003; Hanski and Gaggiotti, 2004),
there is unequivocal evidence that these landscape-altering processes pose a significant threat to global terrestrial biodiversity.
Understanding the complex and often interlinked causes and consequences of habitat loss and fragmentation, and determining
effective means to mitigate and prevent its harmful influence, is therefore critical to preserving terrestrial ecosystem functioning and
biodiversity into the future.
Reference Module in Earth Systems and Environmental Sciences https://doi.org/10.1016/B978-0-12-409548-9.10913-3 1

2 Impacts of Habitat Loss and Fragmentation on Terrestrial Biodiversity
Fig. 1 Habitat loss and fragmentation pictured throughout a landscape in the Monteverde Ecoregion of Costa Rica. Photo by J.E. Rogan.
The underlying mechanisms of habitat loss and habitat fragmentation and their potential consequences have been covered
extensively through seminal works by such authors as Lovejoy, Fahrig, Andren, MacArthur and Wilson and many more.
The Theory of Island Biogeography
Concepts of habitat fragmentation emerged from the expansion of theories of island biogeography (MacArthur and Wilson, 1967).
Hypotheses developed early on based on island biogeography offered a theoretical framework to understand how fragmentation
would influence population viability and persistence in terrestrial landscapes composed of habitat patches, or “islands,” in a “sea”
of human-modified land. The two predictor variables in this theory are island isolation and island size. Based on this theory, one
should expect more species to persist in larger patches and regions with more connectivity amongst patches. However, the concept
of isolation was adapted once this theory became conceptually extended from island systems to terrestrial biomes (Haila, 2002).
Isolation was now described as the result of habitat loss, and it represented the distance from one patch to its neighboring patches
rather than the distance to a mainland. Because of its roots in island biogeography, isolation was viewed as representing habitat
subdivision, a characterization of habitat fragmentation, even though it was inextricably tied to habitat loss. The Island Biogeog-
raphy theory was therefore amongst the earliest theories to predict immigration and extinction and patterns of species richness in
isolated habitats, and later applied to predict the influence of fragmentation on these variables. Experiments in continental settings
then followed in order to test the theory, giving rise to new perspectives.
One clear difference and potential flaw in the application of this theory to terrestrial biomes that scientists began to unravel is
that islands are surrounded by sea, an entirely inhospitable matrix for island-dwelling species. Habitat fragments are distinct from
islands as they can be surrounded by a matrix that may not actually be completely unsuitable for particular species. Thereby, the
influence of terrestrial habitat fragmentation on species dynamics should also be distinct.
SLOSS: Single Large or Several Small?
An early debate in habitat loss and fragmentation research was whether one single large patch of habitat or several small habitat
patches are more effective at conserving biodiversity over the long-term in order to guide the development of reserves that maximize
the benefits for species conservation. The role of reserve size working in conjunction with functional traits of organisms such as
dispersal ability and source-sink dynamics can influence the outcome on species response (Lasky and Keitt, 2013). Suitability has
been found to vary across taxa and landscapes. Beta diversity was found to be higher amongst many small patches in an
experimental landscape than patches that were larger but arranged in a similar fashion (Cook et al., 2005). Tscharntke et al.
(2002) found that 10 ha of 29 small grassland patches of protected area held many more species than a few larger habitat patches of
the same area. However, there is now a substantial level of evidence to support the notion that large, continuous tracts of habitat are
generally more ideal for long-term species population persistence (Laurance et al., 2012; Ewers and Didham, 2006).
Defining and Distinguishing Habitat Loss From Habitat Fragmentation
Habitat fragmentation is commonly described in the literature as the consequent result of habitat loss in which large, continuous
habitat is broken up into many smaller fragments with less overall area that are separated from one another by a human-modified
matrix of different land use types (modified from Didham, 2010; Ewers and Didham, 2006). This definition incorporates
components of land use change related to both the total amount of remnant habitat as well as its spatial configuration within the
same all-encompassing phrase. Defining habitat fragmentation and habitat loss, however, is a common point of contention in the
literature as other authors argue that the term habitat fragmentation should refer only to a narrower set of habitat characteristics.
While acknowledging that habitat fragmentation, like habitat loss, is a landscape scale process, these authors assert that these
processes, and their effects on biodiversity, should be independently recognized and assessed (Fahrig, 2003). As influences on
species resulting from altered spatial configuration increase markedly with reduction in overall habitat amount in a landscape,
Fahrig (2003) and other authors have contended that the influence of the remnant habitat amount must first be taken into
consideration before the effects of habitat fragmentation can be independently defined. Thereby, these authors believe that habitat
fragmentation itself has little effect on species and ecosystem dynamics in fragmented landscapes, and that habitat loss instead is
primarily responsible, regardless of its spatial configuration, for the dynamics that follow.
Habitat fragmentation simply is used to describe the way in which remnant habitat patches are distributed after habitat is lost.
Many early studies may certainly have been flawed in ascribing all effects on species in fragmented landscapes to spatial
configuration without considering the correlated nature of configuration with other processes related to overall habitat amount
in the landscape (Fahrig, 2003). However, the principal conclusion of Fahrig (2003) and others (Collingham and Huntley, 2000;
Fahrig, 1997; Flather and Bevers, 2002; Henein et al., 1998), that the impacts of habitat fragmentation on biodiversity is
insignificant after considering habitat amount, is hotly contested and is in in stark contrast to the thousands of studies exhibiting
the pronounced ecological influence imposed by components of spatial configuration such as edge effects, isolation and patch area
after controlling for habitat amount (Thornton et al., 2011a;Haddad et al., 2015). Based on these conflicting views and ways of
approaching experiments on the impacts of habitat loss and fragmentation, it can be difficult to pull out generalizable consequences
of these processes on terrestrial biodiversity across the board. Therefore, some authors believe an all-encompassing “umbrella” term
should be recognized that embodies all of the underlying mechanisms and impacts associated with habitat loss and fragmentation
(Didham, 2010). Habitat fragmentation is arguably still the most frequently used term in the literature to describe the sequence of
interwoven processes occurring in human-modified landscapes, whether or not the direct and indirect underlying relationships
between landscape and patch variables are explicitly acknowledged and distinguished.
Threshold Hypotheses
Attempts to predict general species responses to habitat loss and fragmentation have led to the establishment of ecological threshold
hypotheses. A threshold in ecology is generally described as a tipping point between stable states of a system in which ecological
functioning ceases or becomes disrupted below a critical level of the ecological feature of interest (modified from Andren, 1994). In
terms of the influence of habitat loss and fragmentation on biodiversity, the feature of interest becomes remnant habitat amount, or
vegetation cover. When this threshold is surpassed, a regime shift occurs resulting in significant changes in the nature and extent of
feedbacks of an ecosystem, and in the ecosystem itself (Pardini et al., 2010). These thresholds in the amount of remaining habitat
therefore are intended to determine how land cover in ecosystems can be properly managed to maintain species populations and
ecosystem functioning in the face of land use and land cover change.
Several researchers have attempted to determine such possible thresholds as a means to provide rules-of-thumb that can help
guide management of landscapes (Andren, 1994; Lande, 1987; McIntyre and Hobbs, 1999; With and King, 1999). However,
differing perspectives regarding the relative influence of habitat amount and configuration have impacted how such thresholds
should be defined. The two most commonly recognized threshold theories are the extinction threshold and the habitat fragmen-
tation threshold. The extinction threshold hypothesis, as posited by Fahrig (2003) and others, hones in on the minimum amount of
habitat in a landscape below which a species is not expected to persist. This concept closely follows the ideology that habitat amount
is the most important feature in land cover change that influence species populations, and therefore does not consider possible
fragmentation effects (Fahrig, 2003). In his seminal work, Andren (1994) proposed a modified threshold concept to that of the
extinction threshold known as the habitat fragmentation threshold hypothesis. This theory includes the influence of habitat
configuration into assessment, positing that below a particular amount of remnant habitat, landscape connectivity becomes eroded
as the landscape is broken into several isolated patches and alpha diversity then becomes dependent on the size or isolation of
remaining patches. In other words, in Andren’s model, thresholds occur as a result of increasing influence of fragmentation effects
below some level of habitat amount. Andren proposed this threshold to be at an intermediate level of habitat around 10%–30%
(Andren, 1994) which has been proposed by several other authors (McIntyre and Hobbs, 1999) with some empirical support in the
literature (Pardini et al., 2010; Estavillo et al., 2013; Garmendia et al., 2013).
Some authors have taken such studies a step further to attempt to uncover species-specific threshold responses (Betts et al., 2007)
or to assess regional differences in threshold levels for the same species across different areas (Rhodes et al., 2008). The overarching
applicability of such generalizable thresholds across systems for different species has been brought into question by several authors
(Lindenmayer and Luck, 2005; Huggett, 2005; Rhodes et al., 2008). Still, many point out the potential utility of defining thresholds
for biodiversity conservation when specific considerations have been made, such as an evaluation of the matrix quality within the
landscape (Pardini et al., 2010; Betts et al., 2007).
Main Drivers of Habitat Loss and Fragmentation

Agriculture and Food Production
The expansion and amplification of agricultural activity now rivals the extent of remaining forest cover, occupying 40% of Earth’s
land surface. Nearly 80% of all threatened terrestrial mammal and bird species are imperiled by habitat loss driven by agriculture
(Tilman et al., 2017). These land use practices alone threaten up over 5400 of the world’s species, around 62% of which are listed as
near-threatened or threatened. Asia’s hairy-nosed otter (Lutra sumatrana), Africa’s cheetah (Acinonyx jubatus), and South America’s
South Andean deer (Hippocamelus bisulcus) are amidst over 2300 species disturbed by land use change for aquaculture and livestock.
There are an additional 4600 species currently threatened from land cover change due to agricultural food production, fuel crops,
and fodder including the African wild dog (Lycaon pictus) and the Fresno kangaroo rat (Dipodomys nitratoides) (Maxwell et al., 2016).
Agricultural expansion accounts for the greatest proportion of habitat loss and fragmentation threatening terrestrial biodiversity
(Tilman et al., 2017).
Logging, Deforestation and Degradation

Deforestation from the combined impact of timber extraction and agricultural expansion has contributed to a total reduction of an
estimated 7–11 million square kilometers of forest over the past 300 years. Unsustainable logging is leading to population declines
of over 4000 forest-specialist species, such as the Myanmar snub-nosed monkey (Rhinopithecus strykeri), India’s Nicobar shrew
(Crocidura nicobarica), and the Bornean wren-babbler (Ptilocichla leucogrammica), (Maxwell et al., 2016). Forest and habitat
degradation and conversion causes additional destruction by reducing the quality and integrity of native habitat. Some of the
more common mechanisms through which native habitat becomes degraded are conversion of native forest or habitat to
agroforestry, perennial tree crops, or timber plantations (Newbold et al., 2013).
Urban Expansion
As populations grow and globalization increases, urban areas are now rapidly growing and threatening to infringe on the habitat of
terrestrial species around the world. Assessments of Amazonian forests aimed at determining the fate of their health and
sustainability into the future predict that placement of roads and highways will be the single most important component
determining the spatial pattern of forest loss, and habitat degradation and fragmentation into the future (Laurance et al., 2009).
Past the direct effects of forest loss in human-modified landscapes from practices such as agricultural and urban expansion,
remaining forest patches are expected to endure further negative consequences as they grow smaller, become more isolated, and
exhibit a greater portion of edge habitat (Skole and Tucker, 1993).
Impacts of Habitat Loss and Fragmentation on Terrestrial Biodiversity

Habitat Loss
Habitat loss has significant, consistently negative effects on biodiversity. Habitat loss negatively influences biodiversity directly
through its impact on species abundance, genetic diversity, species richness, species distribution, and also indirectly. For example,
habitat loss has been shown to decrease population growth rate, which was supported by findings from Donovan and Flather
(2002) who demonstrated that species with declining trends in abundance globally had a higher probability of occurring in regions
with high levels of habitat loss than those exhibiting stable or increasing trends. Habitat loss has also been found to lessen the
number of large, specialist species, disrupt species interactions, reduce trophic chain length, diminish dispersal ability and breeding
success, and alter predation rate, and components of animal behavior related to success rates in foraging (Fahrig, 2003).
There are additional important effects that changes in spatial configuration due to fragmentation of remaining habitat has,
however, which act independently of strict habitat loss (Tscharntke et al., 2002; Fahrig, 2003).
Habitat Fragmentation
The greatest driver of fragmentation operating over large spatial scales is anthropogenic habitat modification (Ewers and Didham,
2005). Many landscapes may be naturally mosaicked, or have natural habitat heterogeneity. However, in these instances, terrestrial
species have had the opportunity to adapt to these natural levels of patchiness over long evolutionary timescales and therefore are
not negatively affected. Habitat fragmentation within human-modified landscapes, on the other hand, are the result of people
altering habitat at unprecedented rates as compared to Earth’s natural evolutionary history (Skole and Tucker, 1993). These rates
substantially exceed the capacity for the majority of species to adapt and persist in the face of rapidly decreasing suitable habitat, and
ever-increasing habitat patchiness (Pimm et al., 1995; Myers and Knoll, 2001).
As a landscape becomes progressively fragmented, a greater number of fragments of varying sizes and shapes are created (Baskent
and Jordan, 1995) which are scattered amongst a matrix of altered habitat (Opdam and Wiens, 2002). The components of a habitat
patch that are influenced by external factors are known as edge habitat, while the inner portions that are not influenced by the
surrounding landscape in such a way are known as core habitat. The component of a fragment that is known as the core habitat is
governed by a complex interaction of fragment shape and size as well as the quality of the matrix habitat surrounding it (Laurence
and Yensen, 1991; Baskent and Jordan, 1995). Fragmentation is not a process that occurs only at the patch-level, though this is the
scale at which the majority of its ecological impacts are detected. Fragmentation can actually only occur when habitat loss at the
landscape-scale results in reduced overall connectivity (Opdam and Wiens, 2002) and the spatial arrangement and size of
remaining habitat is impacted (Baskent and Jordan, 1995).
Thousands of studies in recent literature have attempted to synthesize the ecological impacts of habitat loss and fragmentation
across landscapes (Fahrig, 2003; Prugh et al., 2008; Thornton et al., 2011b; Tscharntke and Brandl, 2004) or have honed in on one
aspect of habitat fragmentation in particular, such as construction of habitat barriers such as roads (Ries et al., 2004). These studies
have demonstrated indisputable evidence for the influence of patch shape, patch area, edge effects, landscape matrix quality and
isolation on community and population level dynamics in human-modified landscapes. However, there are still incongruences
amongst the results of studies focusing on differing taxa and ecosystems.
Influence of Habitat Fragmentation

Habitat fragmentation typically coincides with overall habitat loss; however, there are at least three distinct mechanisms that
mediate the ecological consequences resulting from fragmentation. First, there are mechanisms directly related to shifts in the spatial
configuration of habitat in the landscape, such as isolation between habitat patches. Second, there are those attributable directly to
the decline in overall habitat area. Lastly, there are those explained by indirect effects due to the interaction of habitat loss and
changes in spatial configuration, and spillover effects or impacts due to the interaction of fragments with the matrix habitat
(Didham et al., 2012) (Fig. 2).
Habitat area
Alongside perhaps that of edge effects, habitat area or patch size and its influence on species occurrence is the most widely studied
variable in fragmentation literature, owing largely to the volume of research dedicated to understanding species-area relationships
(Lomolino, 2000). Species-area relationships are a common focus of habitat fragmentation research (Seabloom et al., 2002; Ewers
and Didham, 2006), though they tend to be a controversial subject area due to complex interactions and conflicting findings
(Simberloff, 1992). The direct reduction of habitat area has been cited as one of the main drivers of species extinctions (Tilman et al.,
1994) and has a characteristically strong, negative effect on biodiversity (Fahrig, 2003). When the overall amount of habitat in a
landscape is reduced, this typically results in a heavily skewed distribution of remaining patches, with several small patches, fewer
large fragments (Ranta et al., 1998; Ewers and Didham, 2007) and increased isolation amongst fragments (Andren, 1994).
Consequently, this contributes to reduced colonization rates and population sizes, increasing the risk of local extinctions (Bowers
and Matter, 1997; Bender et al., 1998; Hanski, 1998; Crooks et al., 2001; Hames et al., 2001; Schoereder et al., 2004). Individual
species have minimum patch size requirements (Diaz et al., 2000). Small patches typically contain fewer species than large patches
(Debinski and Holt, 2000), and the species within smaller patches typically represent a subset of the species found within larger
patches (Ganzhorn and Eisenbeiß, 2001; Kolozsvary and Swihart, 1999; Vallan, 2000). The amount of habitat on a landscape
required for species occurrence similarly varies (Gibbs, 1998; Vance et al., 2003). Landscapes with less habitat should therefore also
contain a portion of the species found in similar landscapes that contain more habitat.
Reduction in habitat area due to habitat loss and fragmentation can have negative influences on species at the community and
population levels. At the community level, there are commonly pronounced shifts in species composition as a result of reduced
patch area (Saunders et al., 1991; Ewers and Didham, 2006), primarily because different species have assorted area and resource
requirements, as well as differential dispersal capabilities. Species such as ground beetles, for instance, have high dispersal abilities
and thus are less affected by decreased area than groups with lesser dispersal capabilities (Didham, 2010). High dispersal rates
contribute to the “rescue effect” by which small populations are able to persist due to the continuous flow of new individuals
between patches. Otherwise, these populations would experience high rates of extinction due to small patch size with no species
Fig. 2 Observations of common community-level species responses to habitat fragmentation. General predictions of how species richness typically changes in
response to the five principle components of spatial alteration as a result of habitat fragmentation. Adapted from Didham, R. (2010). Ecological consequences
of habitat fragmentation. In: Jansson, R. (ed.) Encyclopedia of life sciences, pp. 1–11. Chichester: John Wiley & Sons, Ltd.
turnover (Didham, 2010). Metacommunity ecology has played a large role in the increased focus on spatial ecology research in
recent years, addressing species interactions within the spatial domain of metapopulations (Leibold et al., 2004; Gonzalez, 2009).
Increasingly it is recognized as a theoretical framework for studying the influence of habitat fragmentation on communities. As there
are frequent interactions amongst species with varied responses to patch area through mechanisms such as competition, it is
difficult to determine how communities are expected to respond to fragmentation when examining metapopulation models of
individual species.
At the population level, the influence of altered spatial configuration of habitat through fragmentation on the abundance and
distribution of individual organisms underpins the theoretical framework of metapopulation ecology (Hanski, 1998). When patch
size decreases, reproductive success of the population decreases, colonization rates are reduced, resources become limited, and
ultimately, the maximum population size is inherently restricted. Natural disasters, decreased genetic diversity, and demographic
and environmental stochasticity are all components that underlie this relationship between patch area and its resulting influences
on populations (Hanski and Gaggiotti, 2004). “Extinction vortices” can result when these mechanisms work in conjunction (Gilpin
and Soulé, 1986). In extreme instances, this places populations at an increased risk of local extinction (Hanski and
Ovaskainen, 2000).
The expected net result of altered community and population dynamics is an overall loss of species richness amongst small
habitat patches (Ewers and Didham, 2006). The positive species-area relationship may be attributed to several factors that have a
strong impact on species in small patches, such as increased disturbance, reduced habitat heterogeneity, reduced resource
availability, or disrupted extinction and colonization rates. However, consequences are in fact context-dependent and are often
mediated by external forces operating on habitat patches such as quality of the surrounding landscape matrix (Didham, 2010).
Reduction in habitat area resulting from loss and fragmentation are commonly used to predict alterations in species diversity.
Any alterations in species richness at the patch scale are best considered as a net result of all the community-level and population-
level changes working in tandem to modify species occupancy across fragmented landscapes (Didham, 2010). Additionally, below
threshold levels of remnant habitat in a region experiencing habitat loss, species and population responses have been shown to be
nonlinear and controlled by both the spatial configuration and the size of the remaining habitat patches (With and King, 2001).
Empirical research investigating fragmentation effects on terrestrial biodiversity, however, has not always found a strong
influence of patch area on species persistence. Metaanalysis of hundreds of terrestrial species in fragmented landscapes across the
globe performed by Prugh et al. (2008) found that birds and mammals were most sensitive to area while amphibians were least
sensitive, and carnivores (including insectivores) were more sensitive than omnivores. The groups with particular sensitivity to
habitat area were also those with higher metabolic requirements rather than those with greater habitat specialization. Arboreal
species were the one area-sensitive group with evident habitat specialization. However, the authors explain that the difficulty in
discerning and defining suitable habitat across studies may explain the low area sensitivity of amphibians in part, as amphibians use
distinct habitats during different life stages and therefore delineating patches may actually underestimate the extent of habitat they
use (Prugh et al., 2008). Overall, they concluded that patch area was often a poor predictor of occupancy. They did, however, find
that the majority of species under study had a higher chance of occurring in less isolated, larger patches and some species were
particularly sensitive to isolation and patch size (Prugh et al., 2008). Similar confounding findings have been shown in other
studies. In multiple studies examining habitat area and butterflies, it was shown that more species overall were found across several
small habitat fragments than in multiple large fragments with the same overall area in grasslands in Germany (Tscharntke et al.,
2002) as well as forests in Spain (Baz and Garcia-Boyero, 1996). These results probably occur because a series of small fragments
spread over a wide geographic area encompass a broader range of environmental heterogeneity than does a single large fragment
(Tscharntke et al., 2002). For long-term persistence, large habitat areas are still considered to be favorable to small areas for species
that are extinction-prone and area-sensitive (Burkey, 1995; Ferraz et al., 2003), as they support larger populations and are more
likely to resist degradation throughout time (Connor et al., 2000; Tscharntke et al., 2002).
Beginning in 1979, the Biological Dynamics of Forest Fragments Project (BDFFP) is now the world’s longest-running and largest
empirical investigation of the effects of fragmentation on biota in its evaluation of organisms in the Amazon rainforest (Lovejoy
et al., 1986; Bierregaard et al., 1992; Laurance et al., 2002; Gardner et al., 2009; Peres et al., 2010). Although fragment sizes in this
project range from 1 to 100 ha, it has vastly increased our understanding of fragment-area effects as a primary objective of the project
(Lovejoy and Oren, 1981; Lovejoy et al., 1984, 1986). Evaluating across many taxonomic plant and animal groups has shown that
species richness of many organisms is reduced with declining fragment area despite constant sampling effort across every fragment.
These declines were evident in tree seedlings (Benitez-Malvido and Martinez-Ramos, 2003), palms (Scariot, 1998), leaf bryophytes
(Zartman, 2003), larger herbivorous mammals, primates (Gilbert and Setz, 2001; Boyle and Smith, 2010), understory insectivorous
birds (Stratford and Stouffer, 1999; Ferraz et al., 2007) and more. Smaller fragments are often insufficient for the long-term
persistence of these species populations (Laurance et al., 2011). A select number of groups, including the ant-defended plants and
their ant mutualists, have not exhibited a significant reduction in diversity with fragment area. Fragment size has also been shown to
influence the rate of species loss, as smaller fragments lose species more rapidly (Lovejoy et al., 1986), particularly when the
surrounding matrix is unable to be traversed or colonized. Ferraz et al. (2003) predicted from these findings for birds that a 1000-
fold increase in fragment area would be necessary to decrease the rate of local species extinctions by 10-fold. A fragment with an area
as large as 10,000 ha is still predicted to experience a marked loss a bird fauna over 100 years (Ferraz et al., 2003). Mark-recapture
data also suggests that very large fragments are necessary to sustain unaltered species assemblages of some groups of birds such as
ant-following birds, which require large forests areas for foraging (Van Houtan et al., 2007) (Fig. 3).
Fig. 3 Map displaying the study area for the Biological Dynamics of Forest Fragments Project (BDFFP) located in central Amazonia. Areas that are unshaded
represent predominantly undamaged forest regions. Adapted from Laurance, W. F., Camargo, J. L., Luizão, R. C., Laurance, S. G., Pimm, S. L., Bruna, E. M.,
Stouffer, P. C., Williamson, G. B., Benítez-Malvido, J., Vasconcelos, H. L., Van Houtan, K. S., Zartman, C. E., Boyle, S. A., Didham, R. K., Andrade, A. and
Lovejoy, T. E. (2011). The fate of Amazonian forest fragments: A 32-year investigation. Biological Conservation 144(1), 56–67.
Fragment shape
Fragment shape and shape complexity is a lesser-studied component of fragmentation research. The more complex the shape of a
habitat patch, the greater level of edge versus core habitat the fragment contains. This in turn has two primary detrimental influences
on species. The first consequence is that a more complex shape can increase the extent to which the patch edge infringes on the core
habitat of the patch, thereby reducing the amount of core habitat available for habitat specialist species to occupy (Collinge, 1996),
This may be particularly apparent and problematic in longer, thinner habitat areas such as those surrounding riparian zones. This is
also an unexpected issue within large reserves. Recent empirical evidence in fragmented landscapes suggests that patch complexity
exponentially increases with increases in patch area, resulting in large patches, such as some large reserves, containing less core and
more edge habitat than previously thought (Ewers and Didham, 2007). These conditions can be further worsened in very large
patches that have been observed to undergo habitat division into several components of core habitat that are isolated from one
another by their surrounding edge habitats (Ewers and Didham, 2007). These disjoint core habitat areas imbedded in large habitat
patches have been found in literature reviews on density-area relationships to stunt species populations to one fifth of what their
potential population size would be within uninterrupted core habitat (Connor et al., 2000). While this assumption needs further
evidence, these processes together indicate that higher shape complexity in habitat patches coincides with greater species loss.
A second primary ecological consequence of patch shape complexity is that greater complexity may lead to both higher patch
colonization rates and emigration rates between patches. This can result in reduced species population persistence and increased
irregularity in population size across patches (Hamazaki, 1996; Collinge and Palmer, 2002; Cumming, 2002).
Matrix influence
Matrix habitat has additional important influences that are worth discussion. Matrix habitat simply refers to human-modified land
that surrounds or intersperses throughout remnant native habitat patches in fragmented landscapes (Didham, 2010). The
relationship between habitat patches and the matrix, however, is complex, primarily due to variability in the quality and structure
of different matrix habitats, and differential responses by individual species to the matrix. Applying the theory of island biogeog-
raphy to terrestrial biomes, falls short when attempting to uncover the influence of fragmentation effects on terrestrial biodiversity,
as the matrix habitat is not an inhospitable ocean, but varies in its level of suitability for different study species. Several species may
be able to successfully exploit resources found within the matrix that can supplement resources that may be found in smaller
quantities in remnant patches (Ries et al., 2004). In these instances, the division between a habitat patch and the human-modified
matrix may be rather fluid, as some species are able to make use of the entire landscape equally. This was found to be the case with
ants that were shown to feed in Mexican coffee plantations surrounding their native forest habitat, and in some cases even live
indefinitely in these areas (Perfecto and Vandermeer, 2002). The degree of habitat connectivity is thereby partially governed by
habitat continuity, but is also the level to which an area of a landscape enables or inhibits the dispersal of individuals between
patches (Langlois et al., 2001). The location of habitat fragments as well as features and quality of the surrounding matrix thereby
determine species movement and colonization in conjunction (Baskent and Jordan, 1995; Ewers and Didham, 2006). Throughout a
human-modified landscape, it is still an expectation that the quality of habitat will differ in relation to the distribution of the
primary land uses present. Therefore, regions that are dominated by land uses that are deemed as being higher quality for species
persistence are more likely to support a higher number of organisms (Perfecto and Vandermeer, 2002).
The number of studies supporting the notion that matrix quality exhibits a strong influence on the composition and abundance
of organisms in habitat patches has increased dramatically in recent years. (Gascon et al., 1999; Cook et al., 2002; Prugh et al., 2008;
Pardini et al., 2010; Thornton et al., 2011a). In a seminal metaanalysis of habitat area and isolation effects across 89 studies of
terrestrial fauna, Prugh et al. (2008) posited that the characteristics of intervening areas between fragments should be of primary
interest, as landscape context was shown to have a strong influence on the patterns of occurrence for numerous taxa and ecosystems
worldwide. Without considering landscape context, another metric, such as patch size, may be falsely cited as the cause of a
particular species response. Fahrig (2003) describes this important distinction with a study by Donovan et al. (1995) on forest birds
where the authors found that these birds reproduced less in smaller patches than in larger patches. However, if the smaller patches
occur in less forested areas, then the lower reproductive rate may not be due to patch size, but rather that there may be more brood
parasites and nest predators that reside in these more open surrounding habitats (Hartley and Hunter, 1998; Robinson et al., 1995).
The conditions in the matrix surrounding fragments can also define the degree to which external ecological conditions penetrate
a habitat patch (Baskent and Jordan, 1995). Characteristics of the matrix have the capacity to influence the movement and dispersal
of organisms from patch to patch. This is mediated by the level of contrast between the structure of the patch and the matrix that
determine how permeable the habitat edges are to movement (Collinge and Palmer, 2002). In conjunction, these components are
often the main components driving colonization-extinction dynamics and biodiversity loss (Brotons et al., 2003; Kupfer et al.,
2006). As a result, many researchers now recognize and emphasize the importance of considering the matrix habitat and matrix
quality when evaluating both patch and landscape components influencing species populations in habitat fragmentation research.
This includes the utility of the matrix habitat to support species outside of protected areas long-term. Restoration of areas of the
matrix to enable species dispersal and population persistence for many organisms at both the patch and landscape scale is now
being promoted as a feasible approach to conservation management in human-modified landscapes (Donald and Evans, 2006).
In the BDFFP, the matrix surrounding habitat patches was found to strongly influence plant communities by altering the
movements of pollinators and seed dispersers (Dick, 2001; Dick et al., 2003; Bobrowiec and Gribel, 2010; Boyle and Smith, 2010),
mediating particular edge effects, and substantially impacting the seed rain that reaches patches (Laurance et al., 2011). Pioneer trees
that regenerated in patches, for example, varied dramatically in configuration from patches that were bordered by Vismia-dominated
regrowth, to those surrounded by regrowth dominated by Cecropia (Nascimento et al., 2006). In such instances, it is possible that
animal and plant communities may develop in such a way that they will begin to reflect the composition of the neighboring matrix
habitat to some degree (Laurance et al., 2006). This has already been documented in areas of the tropics (Janzen, 1983).
Edge effects
As habitat is lost and habitat patches decrease in size, the extent of edge habitat in a landscape begins to dramatically increase,
ultimately making interior core habitat vulnerable to outside influences (Didham, 1997). Edge effects can be defined as ecological
alterations linked with development of sudden, artificial edges of forest fragments (Didham et al., 1998). These influences occur
between two bordering land-use types where a shift in biotic and abiotic variables takes place (Cadenasso et al., 2003). As compared
to interior habitat, edge habitat is characteristically drier, windier and hotter, and experiences altered habitat structure and plant
composition, and greater light exposure (Matlack, 1994; Williams-Linera et al., 1998; Chen et al., 1999; Harper et al., 2005). As a
result, the abundance of individuals and their patterns of habitat use are often also substantially altered (Ries et al., 2004).
Significant increases in the turnover rate of species composition, and a characteristic rise in species richness, particularly for certain
invertebrates, have been frequently documented at habitat edges (Didham, 1997; Ewers et al., 2007; Ewers and Didham, 2008).
However, the overall changes that occur in species composition and richness at edges are a result of the combined responses from
individual species, which exhibit substantial variation amongst species within a study, and across multiple studies (Didham et al.,
1998; Davies et al., 2000). In general, habitat specialists are expected to decrease in number at habitat edges. Therefore, increases in
species richness are primarily attributed to an inflow of species from surrounding matrix habitat (Didham, 2010). However, it
should be noted that the interpretation of edge effects has largely been drawn from studies that focus exclusively on impacts that
occur within the patch, rather than expanding analysis to also consider external influences where the majority of the drivers of edge
effects occur in the first place (Fonseca and Joner, 2007; Ewers and Didham, 2008). Taking this one-sided view of edge effects
therefore may limit what we can garner about its overall influence on species. Furthermore, responses of some edge variables may
not even be apparent until one observes far beyond the structural edge of vegetation on a particular patch, causing such factors to be
completely overlooked if sampling occurs only within the patch boundaries. Such may be the case with variables that have more
complex properties such as wind turbulence, shading and spillover effects (Cadenasso et al., 1997). A more comprehensive
assessment of edge effects would be a “two-sided” approach, emphasizing the three-dimensional nature of edges as areas of
transition between habitat types that lack distinctly measurable dimensions, unless comparisons are made in relation to the interior
matrix habitat as well as the neighboring patch (Cadenasso et al., 2003). In edge effect studies, some authors have also emphasized
the importance of distinguishing between edge extent, described as the distance between the patch and the matrix habitat where a
statistical difference in response can be identified, and the edge magnitude, or the level of response variation from the interior of the
matrix habitat to that of the patch (Ewers and Didham, 2007). As these components have been found to have unique impacts, it is
important to explicitly distinguish between the two when measuring edge effects (Harper et al., 2005; Ewers and Didham, 2006).
Many studies have focused primarily on edge extent rather than magnitude, though the ecological consequences of edge habitat vary
substantially depending on whether the edge extent or magnitude is greater. An edge response with a large edge magnitude but
comparably small extent may be recognized as more harmful than that with a greater extent and smaller magnitude.
Overall, edge effects are difficult to properly quantify and generalize for several reasons, especially because edge extent is
particularly inconsistent between various edge types and different response variables (Laurance et al., 2002). Many studies also fall
victim to common shortcoming in study design that make results unreliable, such as lacking necessary statistical precision to
successfully define edge extent (Ewers and Didham, 2006) and taking a one-sided approach to quantifying edge effects (Fonseca and
Joner, 2007). With its highly variable nature, edge extent is highly dependent on the response variable being measured and can vary
from just a few meters all the way to a kilometer or more (Ewers and Didham, 2008). Even with several thousands of studies
examining edge effects, there remains little understanding of the elements that define edge extent, or discriminate between edge
magnitude and extent to begin with (Didham, 2010).
Laurance et al. (2002) concluded in an assessment of the Biological Dynamics of Forest Fragments Project (BDFFP) that in
Brazilian tropical forest fragments edge effects are one of the greatest drivers of ecological modifications in these forests, and exhibit
strong negative effects on several taxa. These effects are apparently much stronger than negative edge effects in temperate systems
(Kremsater and Bunnell, 1999). Edge effects have been shown to be extremely variable in their impact including influences such as
elevated windshear, desiccation stress, and wind turbulence that drastically increases levels of tree damage and death (Laurance
et al., 1997, 1998). These changes have a domino effect, leading to far-reaching modifications in the community composition of
plants, particularly lianas (Laurance et al., 2001) and trees (Laurance et al., 2000, 2006). These pressures have the capacity to restrict
the formation of shade-tolerant plant species in fragments (Uriarte et al., 2010) and reduce germination (Bruna, 1999), resulting in
drastic alterations in the number and composition of tree seedlings (Benitez-Malvido, 1998; Benitez-Malvido and Martinez-Ramos,
2003). In addition, research offers strong support for the notion that edge effects can become cumulative when there are several
edges in close proximity, making for more dire edge impacts than a single effect acting independently. This conclusion is
additionally reinforced by studies of edge-related changes in tree mortality (Laurance et al., 2006), forest microclimate (Kapos,
1989), species richness and abundance of tree seedlings (Benitez-Malvido, 1998; Benitez-Malvido and Martinez-Ramos, 2003),
vegetation structure (Malcolm, 1994), the diversity and concentration of disturbance tolerant pioneer trees (Laurance et al., 2006)
and liana abundance (Langlois et al., 2001). These additive effects of neighboring edges may also elucidate why complexly shaped
or small (<10 ha) fragments frequently undergo drastic modification after fragmentation (Zartman, 2003; Laurance et al., 2006).
Many taxa, including several flies, bees, wasps (Fowler et al., 1993), ants (Carvalho and Vasconcelos, 1999), butterflies (Brown
and Hutchings, 1997), understory birds (Laurance, 2004), and beetles (Didham et al., 1998), become reduced in number by forest
fragment edges. Along forest roads 20–30 m in diameter, there are even obvious negative edge effects that take place in large tracts of
forest. For example, within understory birds, five out of eight foraging guilds were found to decrease substantially in number within
70 m of roads, with associated reduction in canopy cover and increased tree mortality (Laurance, 2004). However, there are some
taxa that either become more abundant or remain neutrally influenced by edge habitat if they have general habitat requirements or
are able to tolerate the surrounding matrix (Pardini et al., 2010). Many frog species (Gascon, 1993), leaf bryophytes (Zartman and
Nascimento, 2006), and wandering spiders (Mestre and Gasnier, 2008), to name a few, are not significantly impacted by edges. On
the other hand, in some cases, hyperabundance in species may occur where particular species thrive at edges or within disturbed
patches. Disturbance tolerant organisms such as butterflies preferring plentiful light and rapid-growing lianas (Laurance and
Williamson, 2001) numerous frugivorous and gap-favoring birds (Laurance, 2004), and hummingbirds (Stouffer and Bierregaard,
1995) are examples of species that actually increase in numbers in edge habitat, sometimes considerably.
Another unique facet of fragments, particularly in edge habitat, is that community and population dynamics and species
interactions are highly variable as compared to those in intact, interior forest habitats. Species interactions at habitat edges become
less stable, which may be partially owing to the fact that edge species are often restricted to only some edge areas of a habitat patch
rather than all of them resulting in higher beta diversity in patch edges versus patch interiors (Didham et al., 1998). Variation in the
abundance of invertebrate predators was shown to decrease from edge to interior habitat within an apple orchard in one particular
study (Brown and Lightner, 1997). Hyperdynamism, described by Laurance (2002) as the rise in the frequency and/or amplitude of
population, landscape and community dynamics in fragmented habitats, often occurs in conjunction with this variability in
interaction. This is primarily because habitat patches are more likely to be influenced by stochaistic events and infiltration of
external processes in the matrix, causing extreme fluctuations or destabilizations in communities of organisms. It is particularly the
case where there are high rates of disturbance and low levels of immigration between patches (Hubbell, 2001). Many examples of
hyperdynamism have been observed in fragmented landscapes. In the BDFFP, butterflies are one example of taxa that have been
shown to be extremely hyperdynamic in their community characteristics, undergoing drastic proliferations in numbers in response
to abrupt increases in their preferred host plant species along habitat edges (Brown and Hutchings, 1997). Bat communities were
also shown to have an uncharacteristically fast turnover of species, likely resulting from large losses of forest-interior species, paired
with an increase of opportunistic frugivores feeding in edge habitat and in nearby areas of regrowth (Laurance et al., 2002). Large-
scale analysis has revealed that almost 20% of the world’s remaining forest is within 100 m of an edge and over 70% of these forests
are within 1 km of a forest edge. Therefore, the majority of Earth’s forests are well within the boundary whereby potentially negative
external influences can harm forest ecosystems (Broadbent et al., 2008).
Isolation
Isolation amongst habitat patches and its antithesis, patch connectivity, cannot be directly quantified as they exist in varying
gradations throughout space and time, and their influence is highly dependent on the individual dispersal abilities of different
species. Greater levels of habitat loss in a landscape cause the degree of isolation and time since isolation to grow in tandem. As a
result, it can be difficult to discern between their individual influences on organisms (Saunders et al., 1991). Directly following
isolation, it is common for a patch to harbor a far greater number of species than it is capable of supporting as viable populations
over the long term. Over time, through the process of “species relaxation,” species richness will become reduced (Brooks et al.,
1999). The amount of time over which relaxation is expected is highly dependent on the average lifespan of individual species and
the initial population size in the patch. Therefore, it can vary anywhere from just a few months, all the way to hundreds of years for
some tree species (Vellend et al., 2006).
Time since isolation can also have a large influence on the ability to make comparisons between species composition and
diversity across patches, as patches that have been isolated longer than others will exhibit a different influence than those more
recently isolated. The influence of distance of isolation and time since isolation are closely linked. While species populations may be
supported over a long period of time if there is only a small distance between patches allowing for the safe passage of some
organisms, a larger distance of isolation may alternatively pose a greater threat to long term species persistence (Didham, 2010). The
ability for an organism to traverse a gap or matrix habitat, however, varies greatly amongst species. For example, the BDFFP revealed
that many organisms in the Amazon would not enter clearings, some of which were even unable to pass over roads through the
forest. Insectivorous understory birds, for instance, were found to have substantially reduced movements across roads of 20–40 m,
and markedly decreased abundance within close proximity to these roads (Laurance et al., 2004). When translocated for experi-
mental purposes, some resident adult birds have been shown to avoid crossing patches of pasture (250 m) but traverse a 50–75 m
highway without trouble in order to reach their home territory (Laurance and Gomez, 2005). Even very small openings of
100–200 m can pose a significant barrier to many taxa that are forest specialists, from epiphyllous lichen spores (Zartman and
Nascimento, 2006; Zartman and Shaw, 2006), dung beetles (Noss, 2008), euglossine bees (Powell and Powell, 1987), to
herbivorous insects (Fáveri et al., 2008). Other sensitive species, however, have still been found to cross forest clearings to move
from smaller fragments to large swaths of forest (Van Houtan et al., 2007). Forest openings that have occurred due to habitat
fragmentation may also threaten the dynamics of the fragment interior by introducing nonforest, foreign species (Gascon et al.,
1999; Laurance et al., 2009).
Due to variation in the degree to which particular matrix habitats inhibit species dispersal (Roland et al., 2000; Ricketts, 2001),
and the differential abilities of various species to traverse the matrix between habitat patches (Haddad and Baum, 1999; Collinge,
2000), there is substantial disparity in the literature as to how isolation is expected to impact species populations (Ewers and
Didham, 2006). This may be help explain why Prugh et al. (2008) found that isolation was a poor indicator of species occurrence in
their analysis of hundreds of terrestrial species in fragmented landscapes. What can be drawn from isolation studies, though, is that
the impact of spatial isolation can be just as dependent on the level of permeability of the surrounding matrix as the configuration
of the habitat edge, and that individual species’ ability to traverse matrix habitat can be equally mediated by behavioral
characteristics as by physical traits that allow them to travel across greater distances (Didham, 2010) (Fig. 4).
Confounding Factors in Drawing Generalizations on the Influence of Habitat Loss and Fragmentation
The most critical recent advances in our understanding of the ecological effects of habitat loss and fragmentation on terrestrial
biodiversity all begin with the acknowledgement of the strong context dependence of ecosystem responses. These notably include:
time lags in species loss, spatial context dependence across several scales, synergistic interactions between habitat fragmentation and
other factors of widespread ecological change and variation in responses mediated by species-specific traits (Didham, 2010).
Spatial Scale of Analysis

The scale of analysis in empirical studies is also a crucially important component in accurately measuring the influence of habitat
loss and fragmentation. Many studies that focus on evaluating the influence of habitat loss and fragmentation have done so at the
patch level scale, rather than the landscape scale. As fragmentation is a landscape-scale process, accurate measurement of
fragmentation and its influence on species biodiversity should also be measured at the landscape scale in order to ensure accurate
evaluation and derive applicable predictions (Fahrig, 2003; McGarigal and Cushman, 2002). An additional issue involving scale in
experimental studies is that the scale of a landscape depends specifically on the study species being evaluated. Therefore,
determining the proper scale to study landscape effects on different taxonomic groups is a critical but often poorly accounted for
component in these studies. In a metaanalysis of 134 fragmentation studies, McGarigal and Cushman (2002) found that only 5%
considered fragmentation from an explicitly organism-relevant scale.
Nonlinearity in Species Response

Species of different taxonomic groups, or even the same taxonomic group, that harbor different functional traits are not expected to
respond in the same way to the influence of habitat loss and fragmentation. Many studies and metaanalyses of studies across
taxonomic groups have shown differential responses of biodiversity to habitat loss and fragmentation effects dependent on their
habitat requirements and degree of specialization (McGarigal and Cushman, 2002; Ewers and Didham, 2005; Prugh et al., 2008;
Pardini et al., 2010; Thornton et al., 2011b; Estavillo et al., 2013). These differences can lead to seemingly conflicting findings across
studies examining different species and taxonomic groups making it more difficult to pull out generalities. Certain traits have been
identified to increase the vulnerability of species to negative effects of habitat loss and fragmentation such as large body size, rarity,
Fig. 4 Fragmentation effects are shown to trickle down throughout aspects of entire ecosystems in several long-term, large scale fragmentation experiments
under analysis. This figure summarizes the main findings of Haddad et al. (2015) of fragmentation effects on ecosystem processes across several long-term, large-
scale fragmentation experiments. Effects on all principal ecosystem processes at all relevant ecological scales is evaluated. (A) Shows reduced area in three long-
term, large-scale fragmentation experiments (the Biological Dynamics of Forest Fragments Project, Kansas Experiment and Wog Wog) and its influence on
ecosystem dynamics in two experiments (SRS Corridor Experiment and Moss Fragmentation Experiment). (C) Shows effect of increased edge on ecosystem
dynamics across all analyzed experiments. (B) Demonstrates influence of increased isolation on ecosystem dynamics. Dots represent the average effect size for an
ecological process. Effect sizes are statistical, so negative or positive values could represent degrading function. Horizontal show the range of the value when a dot is
signified by multiple studies. Adapted from Haddad, N. M., Brudvig, L. A., Clobert, J., Davies, K. F., Gonzalez, A., Holt, R. D. and Cook, W. M. (2015). Habitat
fragmentation and its lasting impact on Earth’s ecosystems. Science Advances 1(2), e1500052.
high trophic level, low dispersal capability, lower abundance prior to fragmentation, lower reproductive rate, and greater habitat
specialization (Laurance, 1991; Didham et al., 1998; Davies et al., 2000; Tscharntke et al., 2002; Henle et al., 2004). When found
together, these traits increase the vulnerability of species to fragmentation effects. Particularly in landscapes that have been
significantly degraded, interactions of multiple traits together can increase the susceptibility of species to the negative consequences
of habitat loss and fragmentation far beyond what would otherwise be expected from the additive effects of several traits measured
individually. Some species traits also may be expected to have a greater influence depending on the specific spatial context of the
region under study (Henle et al., 2004).
Temperate Versus Tropical Systems

Through comparative evaluation of empirical studies conducted in tropical and temperate ecosystems, a clear distinction in
predicted response to habitat loss and fragmentation has been uncovered. While Fahrig’s (2003) theoretical assumption that effects
of habitat loss outweigh those of fragmentation was supported by some empirical studies in temperate ecosystems, the response was
found to differ when evaluated within tropical biomes. In studies that explicitly controlled for the independent effects of habitat loss
and fragmentation on medium and large tropical mammals, both Estavillo et al. (2013) and Thornton et al. (2011b) found that
these species responded strongly to habitat fragmentation effects independent of habitat loss. Additional findings by Laurance et al.
(2002) indicated strong negative edge effects on several taxa within tropical Brazilian forest that were much stronger than those
found in some temperate systems (Kremsater and Bunnell, 1999). These findings indicate that response of species to habitat loss
and fragmentation not only depends on factors such as the taxonomic groups under evaluation, but also on the geographic location
in which these processes take place.
Synergistic Interactions Between Fragmentation and Other Threatening Processes

Habitat fragmentation does not occur in isolation from other global threats to biodiversity. Synergistic interactions amongst
multiple drivers of biodiversity loss and ecological change may significantly augment the detrimental impacts of both processes,
or mask the independent influence of fragmentation on species (Ewers and Didham, 2005). For example, fragmentation can disrupt
pollination systems, or increase the rate of disease transmission, leaving populations in habitat remnants susceptible to human
encroachment, fire and introduced species, and may amplify the vulnerability of species to climate change. Habitat fragmentation
can create synergies and exacerbate the rate and extent of other processes such as species invasions and climate change (Ewers and
Didham, 2007). The effects of climate change are expected to be much more extreme in human-modified landscapes as the ability
for habitat specialists to undergo range shifts in response to changing environmental conditions will be heavily restricted in
fragmented landscapes, versus those that have not been significantly altered (Thomas et al., 2006).
Time Lags
The effects of habitat loss and fragmentation are expected to differ between the short and the long term. Short-term effects have been
better studied, with most studies investigating influence <100 years after fragmentation (Watson, 2002), and potential long-term
consequences have only begun to come to light in more recent years. Not all species show reactions immediately after habitat
change, indicating that many studies fail to pick up on the real influence of common fragmentation effects. Species also differ in the
length of time it takes for them to respond, further confounding results. Crowding effects, or the short-term influx of population
densities of species that can invade remnant patches, have been widely documented immediately following disturbance (Collinge
and Forman, 1998). This, however, gives an unrealistic picture of the long-term environmental consequences of habitat loss and
fragmentation as this response is transient in nature (Ewers and Didham, 2006). Over the subsequent process of species relaxation,
populations are reduced and different species are lost at different rates depending on individual traits and characteristics of remnant
habitat. This can span from a few months to many years for some species (Brooks et al., 1999). In areas that have undergone
particularly extreme and accelerated habitat loss and disturbance, longer time-lags in species response could lead to what would be
an enormous “extinction debt” where hundreds to thousands of threatened species could suddenly be lost in the next 100 years
(Tilman et al., 1994). The potential for such extinction debts has been supported by a growing number of empirical studies in recent
years as the temporal variation in community response following fragmentation is being steadily unveiled (Ewers and Didham,
2006). The idea of temporal reliance of processes occurring in human-modified landscapes has also been uncovered for theories
such as “colonization credit,” or the number of species expected to later colonize an area following habitat restoration (Cristofoli
et al., 2010) and “invasion debt,” describing the number of species that have not yet occupied a patch from emergent populations
throughout the greater landscape (Seabloom et al., 2006).
Lessons From Large-Scale, Long-Term Fragmentation Experiments
Long-term and large-scale field-based fragmentation experiments are crucial in expanding our understanding of the influence of
habitat fragmentation on terrestrial biodiversity. Experimental approaches guided by the BDFFP in the Brazilian Amazon (Bierre-
gaard Jr. et al., 1992; Laurance et al., 2002), have increased in prevalence (Debinski and Holt, 2000; McGarigal and Cushman,
2002), with projects now ranging from spatial scales as large as the entire Earth, to those as small as microhabitats (Burkey, 1995;
Gonzalez and Chaneton, 2002; (Riitters et al., 2000). The most broadly acknowledged fragmentation experiments encompass a
wide range of ecological communities (Haddad et al., 2015). Unlike strictly observational studies, these projects employ carefully
constructed experimental designs, developed prior to the start of data collection in the field. They include substantial levels of
replication and collect information on conditions prior to disturbance, allowing for more powerful inferences to be made (Wilson
et al., 2016). These studies are also able to isolate the influence of fragmentation from habitat degradation and loss, and control for
the impact of differing quality of the surrounding matrix habitat that confound fragmentation effects by influencing ecosystem
functioning and species composition within patches (Flinn et al., 2005). These studies are therefore a highly influential means of
increasing our understanding about the broad and varied impacts of habitat fragmentation across landscapes (McGarigal and
Cushman, 2002).
In a metaanalysis across all of the largest and longest running habitat fragmentation experiments to date, Haddad et al. (2015)
extrapolated findings to provide insight into some of the most pronounced and far-reaching impacts of habitat fragmentation on
terrestrial biodiversity. These studies, including the BDFFP referenced previously, each account for confounding factors such as
overall habitat amount in the landscape by experimentally forming or removing specific quantities of habitat over replicate
landscapes. They are otherwise widely diverse as they operate across many different ecoregions, and focus on individual aspects
of fragmentation, such as isolation, habitat size and connectivity (Haddad et al., 2015). The largest scale of these studies are
specifically designed with fragment sizes to match those characteristically formed in human-modified landscapes. Together they
cover a wide range of edge distances typically encountered in over half of the earth’s forests (Haddad et al., 2015). Due to their rigid
design, these experiments are highly comparable, allowing for powerful inferences to be made on the underlying components that
drive ecological consequences of fragmentation. The authors explicitly examined the impact of decreased fragment size, elevated
edge habitat and increased fragment isolation on abundance, extinction, dispersal, community composition, ecosystems function-
ing and species richness (Haddad et al., 2015) (Fig. 5).
In their analysis, Haddad et al. (2015) found that fragmentation has some unexpected, pronounced, overwhelmingly consistent,
and primarily negative influences on organisms and local biophysical conditions. From examination of population level responses,
overall patterns in the ecosystem and community reactions were interpreted. Increased isolation between fragments reduced
dispersal and accompanying recolonization rates of patches following local extinction, and smaller patches harbored fewer
Fig. 5 The seven major large-scale, long-term fragmentation experiments across the globe. Arrows represent the beginning of each experiment that has been
ongoing since this time of conception, with the exception of the Moss experiment which is comprised of a series of studies that has been underway for almost
20 years, Components incorporated into the study are denoted with a checkmark. The area shown represents the largest patches in each experiment. The dominant
plant community is listed below each “Matrix” and “Fragment” section for each study, and the relative stage of succession of several experiments. Adapted from
Haddad, N. M., Brudvig, L. A., Clobert, J., Davies, K. F., Gonzalez, A., Holt, R. D. and Cook, W. M. (2015). Habitat fragmentation and its lasting impact on Earth’s
ecosystems. Science Advances 1(2), e1500052.
organisms. While individual species differed in their responses, the abundance of mammals, insects, plants and birds decreased
overall with decreasing patch size and increasing isolation (Haddad et al., 2015). This in turn likely resulted in the reduced capacity
of species to persist in more isolated, smaller patches. Greater edge habitat resulted in higher rates of predation on bird nests thus
reducing bird fecundity, while increased edge habitat in conjunction with reduced patch size and increased isolation lowered
herbivory and seed predation.
In accordance with established theories, species richness of animals and plants across all experiments was significantly reduced
by habitat fragmentation (MacArthur and Wilson, 1967; Tilman et al., 1994; Hanski et al., 1996) with frequent alteration of
composition of whole communities. Within grasslands ecosystems, patch size was found to alter the rate of succession to the extent
that modified seed pools and increased light exposure in smaller patches inhibited the rate of ecological succession compared to
that in larger patches (Cook et al., 2005). In tropical forest biomes, smaller fragment size and greater extent of edge resulted in
modifications in physical characteristics of the environment leading to the loss of older, larger trees in place of pioneer trees. This
had consequent effects on the community composition of insects (Laurance et al., 2000).
Spanning numerous studies and ecosystems across all experiments, fragmentation unfailingly resulted in ecosystem degradation
through the reduction species persistence, trophic dynamics, nutrient retention, dispersal and species richness (Haddad et al., 2015).
Overall, it was clearly demonstrated that factors that may play a small role in the dynamics of fully intact landscapes can come to be
major drivers of change within fragmented regions. This has been shown through such findings as lessened fire frequency in prairie
habitats (Leach and Givnish, 1996) and elevated wind shear in tropical forest patches (Laurance et al., 2011). This analysis also
revealed the long-term consequences of habitat fragmentation in each of the experiments, unveiling that many of the negative
effects on ecosystem functions and species not only persisted, but actually increased in magnitude over time. In the long-term
Kansas Experiment evaluated by Haddad et al. (2015), for instance, plant succession in response to fragmentation was not even
evident until 12 years following fragmentation. These results therefore provide backing for the presence of the ecological processes
of immigration lag, extinction debt and ecosystem function debt as consequences of long-term, gradual impacts of fragmentation
(Tilman et al., 1994; Haddad et al., 2015).
These experiments were thereby shown to play a critical role in both testing current theories and the development of new,
progressive theories. Those important findings have guided the development of more recent theories such as that of the metacom-
munity theory (Leibold and Mikkelson, 2002) that considers components found to be of primary importance in mediating
fragmentation impacts: spatial dynamics, spatial variation in interspecific interactions, variability in matrix habitat quality, and
degree of connectedness amongst patches. Additionally, variation in species-specific traits such as dispersal ability, reproductive
output and trophic level are important in mediating the influence of fragmentation on biodiversity (Haddad et al., 2015).
Accounting for differences in these traits will help to elucidate deviations from the generalizable pattern of overall loss of species
richness associated with habitat loss and fragmentation across taxa and ecotones. These are controlled experiments that account for
specific factors such as patch shape; however, these detected impacts are expected to be even more pronounced and dire in actual
fragmented landscapes.
Conclusions and Future Directions
Comparative findings across several studies focused on impacts of habitat loss and fragmentation at the landscape scale have been
shown to be highly effective in drawing conclusions on the relative influence of increasing patch isolation, patch shape, decreasing
patch area, edge effects, matrix quality and overall habitat loss on different animal communities and terrestrial ecosystems. Such
assessments are critical in allowing researchers to develop more generalizable, overarching consequences of habitat loss and
fragmentation based on unique, system-specific responses (Wilson et al., 2016).
While increasing numbers of these empirical studies in recent years has led to substantial evidence and insight into the potential
consequences of these processes on species population viability and persistence, there remains a dearth of evidence for many
terrestrial taxonomic groups. In metaanalyses of research on the influence of habitat loss and fragmentation on terrestrial
biodiversity, McGarigal and Cushman (2002) and Thornton et al. (2011a) found that birds were the most commonly evaluated
taxa, with mammals and plants following closely behind. Herpetofauna and invertebrates were the least studied taxa, with reptiles
in particular being largely underrepresented (McGarigal and Cushman, 2002; Thornton et al., 2011a). As far as ecosystems,
temperate biomes were the most commonly used study regions by far, followed by tropical systems. Grassland, steppe and desert
were the most highly underrepresented study systems by a significant margin (McGarigal and Cushman, 2002).
Additionally, many pressing questions remain unanswered, including how components of fragmentation interact and influence
other evolutionary processes related to gene flow and alterations in selections patterns, or what some of the more long-term
consequences of fragmentation might be. Conclusions drawn from research to date are also expected to be conservative in their
assessment as the influence of other ecological drivers of change have primarily been omitted. Most forms of global environmental
change found to diminish species populations, such as invasive species, hunting, climate change, and pollution, will be intensified
by fragmentation, and vice versa (Haddad et al., 2015). Over the next 50 years, such threats to biodiversity are only expected to
increase in accordance with rising per capita income and human populations. Areas that have already experienced and continue to
undergo habitat loss and fragmentation, including many deemed conservation hotspots (Myers et al., 2000), are predicted to face
greater risk of extinction for numerous terrestrial taxa (Tilman et al., 2017).
More robust evidence of the impacts of habitat loss and fragmentation on ecosystem functioning and biodiversity, particularly
through complex interactions with other global drivers of change are therefore crucially necessary to accurately grasp the expected
long-term outcome of this pervasive environmental threat. Some such studies are already underway, such as the SAFE Project in the
rainforest of Borneo (Ewers et al., 2011) that consists of a fragmentation experiment embedded within an agricultural plantation
where poaching occurs. Long-term experiments are now in even greater demand in order to acknowledge and plan for anticipated
long-term consequences, and to better understand the relationship between short and long-term effects. The most valuable
forthcoming studies will make good use of emergent tools and technologies like high quality satellite imagery and spatial modeling
with existing ecological theory to increase the understanding of the mechanisms driving habitat loss and fragmentation effects.
Studies that use open access databases of biodiversity and land use data across the globe such as BIOFRAG (Pfeifer et al., 2014),
GBIF (Flemons et al., 2007), Living Planet Index (Loh et al., 2005), and the IUCN Red List (IUCN, 2018), will play a substantial role
in illuminating our understanding of the broad effects of habitat loss and fragmentation on biodiversity into the future. These
findings will be crucial for guiding prioritization and management of land in fragmented regions for conservation and ecological
restoration purposes. Protected areas were once seen as the first line of defense in response to prevalent land use change, now
covering around 14% of Earth’s terrestrial surface (Butchart et al., 2015) with partial success in lessening hunting, extinction risk,
and habitat conversion within their borders (Tilman et al., 2017). However, terrestrial biodiversity continues to decline globally.
While protected areas are still recognized as important, central mechanisms for biodiversity conservation (Watson et al., 2014) they
are also increasingly recognized as inadequate at maintaining numerous taxa over the long-term on their own (Chazdon et al.,
2009). This has been cited for a number of reasons, primarily as they are improperly enforced and guarded in many areas and simply
cannot cover large enough areas including the full home ranges of many species. Over 90% of tropical forests, for instance, exist
outside the confines of reserves and protected areas, leaving a large extent of viable habitat unprotected (Chazdon et al., 2009).
Habitat corridors too have been shown to have mixed results for protection of species, but overall there is an abundance of evidence
from studies across multiple scales that illustrate the potential of corridors to increase species diversity and population abundance
within fragments. They can also substantially influence ecosystem function and species interactions throughout fragmented
landscapes (Villard and Metzger, 2014).
Many now recognize the potential of the matrix habitat to support species populations through more substantial, progressive
management strategies incorporating social and ecological components of human-modified ecosystems (Gardner et al., 2009).
Some matrix types in particular have been shown to be preferable for dispersal of forest species, including those developed through
agroforestry (Villard and Metzger, 2014). Proactive initiatives, such as improved land-use planning, changes in agricultural
practices, and increased conservation-based agricultural trade will also play a large role in the future conservation of species in
complex systems (Tilman et al., 2017).
Terrestrial biodiversity is currently, and will continue to be, severely threatened by anthropogenic drivers of land use change
resulting in more pervasive habitat loss and fragmentation. Coming to an agreement on the approaches and proper terminology for
defining and evaluating these complex and interacting processes in empirical research is an important step allowing for more
general conclusions to be made on their overall impact, and to guide more effective conservation management strategies into
the future.
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Impacts of Habitat Loss and Fragmentation On Terrestrial Biodiversity

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Impacts of Habitat Loss and Fragmentation on Terrestrial Biodiversity

Introduction: Impacts of Habitat Loss and Fragmentation on Terrestrial Biodiversity 1

Introduction: Impacts of Habitat Loss and Fragmentation on Terrestrial Biodiversity

Reference Module in Earth Systems and Environmental Sciences https://doi.org/10.1016/B978-0-12-409548-9.10913-3 1

The Theory of Island Biogeography

SLOSS: Single Large or Several Small?

Defining and Distinguishing Habitat Loss From Habitat Fragmentation

Main Drivers of Habitat Loss and Fragmentation

Logging, Deforestation and Degradation

Impacts of Habitat Loss and Fragmentation on Terrestrial Biodiversity

Influence of Habitat Fragmentation

Spatial Scale of Analysis

Nonlinearity in Species Response

Temperate Versus Tropical Systems

Synergistic Interactions Between Fragmentation and Other Threatening Processes

Lessons From Large-Scale, Long-Term Fragmentation Experiments

Conclusions and Future Directions

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