Biodiversity and Conservation 8: 1383–1397, 1999.

© 1999 Kluwer Academic Publishers. Printed in the Netherlands.

Effects of climate change on biodiversity:

a review and identification of key research issues

MAARTEN KAPPELLE, MARGRET M.I. VAN VUUREN and PIETER BAAS

Rijksherbarium/Hortus Botanicus (RHHB), Leiden University (RUL), P.O. Box 9514, 2300 RA Leiden,
The Netherlands; ∗ Present address: Instituto Nacional de Biodiversidad (INBio), Apartado Postal
22-3100, Santo Domingo de Heredia, Costa Rica (fax: +506-244-2816; e-mail: [email protected])

Received 5 January 1999; accepted in revised form 26 January 1999

Abstract. Current knowledge of effects of climate change on biodiversity is briefly reviewed, and results
are presented of a survey of biological research groups in the Netherlands, aimed at identifying key
research issues in this field. In many areas of the world, biodiversity is being reduced by humankind
through changes in land cover and use, pollution, invasions of exotic species and possibly climate change.
Assessing the impact of climate change on biodiversity is difficult, because changes occur slowly and
effects of climate change interact with other stress factors already imposed on the environment. Research
issues identified by Dutch scientists can be grouped into: (i) spatial and temporal distributions of taxa;
(ii) migration and dispersal potentials of taxa; (iii) genetic diversity and viability of (meta) populations of
species; (iv) physiological tolerance of species; (v) disturbance of functional interactions between species;
and (vi) ecosystem processes. Additional research should be done on direct effects of greenhouse gases,
and on interactions between effects of climate change and habitat fragmentation. There are still many gaps
in our knowledge of effects of climate change on biodiversity. An interdisciplinary research programme
could possibly focus only on one or few of the identified research issues, and should generate input data
for predictive models based on climate change scenarios.

Key words: biodiversity, climate change, research issues

Introduction

Climate change poses a potential threat to the earth’s biodiversity. In comparison to

threats by other human-induced environmental changes (e.g., changes in land cover
and use (Dale 1997), pollution, effects of increased concentrations of greenhouse
gases), direct effects of recent climate change on biodiversity will be slow and diffi-
cult to measure, but the processes are global and practically irreversible. Moreover,
climate change will exacerbate the stresses already imposed on the environment. For
example, in a fragmented landscape, species may be unable to move to a climatic-
ally more favorable environment, because their dispersal capacities are insufficient to
cross the barriers between the remaining natural areas.
1384

In this paper, current knowledge of effects of climate change on biodiversity are

briefly reviewed (Figure 1). In addition, results are presented of a survey of biolo-
gical research groups in the Netherlands (Van Vuuren and Kappelle 1998), aimed at
identifying key research issues for effects of climate change on biodiversity.

Figure 1. Schematic diagram illustrating the alleged human and climatic forces leading to loss of
biodiversity.
 1385

Methods

The methodology of this study consisted of: (i) a review of the bibliography on
the effects of climate change on biodiversity; and (ii) interviews held in 1997 with
ca. 30 biological research groups in the Netherlands, involved in biodiversity and
climate change research. Interviews aimed at identifying key issues in research on
effects of climate change on biodiversity. Results from the interviews were discussed
with participating scientists during a workshop at Leiden University, on 21 November
1997. This workshop contributed strongly in: (a) defining gaps in our knowledge of
effects of climate change on biodiversity; and (b) setting priorities among the identi-
fied key research issues in this field. Scientists who participated in both the interviews
and the workshop suggested a broad range of themes for research. These themes were
used to formulate recommendations for a concise two-year research program to be
carried out within the Dutch National Research Programme on Global Air Pollution
and Climate Change (NRP) at the turn of the millennium.

Results and discussion

Global climate change

Global climate change is one of the most contentious topics in environmentalism,

ecology and politics (Houghton et al. 1990, 1992, 1996; Watson et al. 1996; Zwerver
et al. 1995). Human-induced increases of atmospheric concentrations of gases such as
carbon dioxide, methane, nitrous oxide and chlorofluorocarbons (CFCs) may result
in unparalleled increases in global temperature (Houghton 1995; Bush 1997). This
would happen through an intensification of the so-called ‘greenhouse effect’ i.e. the
absorption of infrared radiation by gases and its re-radiation back toward the surface
of the earth. Measurements over the past 130 years show that atmospheric tempera-
tures have already risen considerably and have been the highest in the last few years.
A drop in global temperatures in 1992 was caused by the eruption of Mount Pinatubo,
a volcano in the Philippines, but this eruption had no lasting effect on trends in global
temperature (data from US Department of Commerce).
In 1996, the Intergovernmental Panel on Climate Change (IPCC) issued its Second
Assessment Report (Watson et al. 1996), which represents the degree of consensus
on various climate change issues. Under the IPCC business-as-usual scenario (i.e.
no reduction in carbon dioxide emissions) global environmental change in the next
century may include an increase of atmospheric carbon dioxide concentrations from
350 ppmv in 1993 to 525 ppmv in 2050. This may imply a 0.3 ◦ C rise in global
mean temperature per decade (Houghton et al. 1992; Sprengers et al. 1994). Recent
climate models calculate a potential rise of the global mean surface temperature of
about 1 to 3.5 ◦ C by 2100 (Watson et al. 1996). Such a change could be 10 to 50
1386

times faster as the natural average rate of temperature change since the end of the last
glaciation (McNeely et al. 1995). However, projected rises in temperature will not
be equally distributed over the globe. Mean temperatures at the poles are expected to
increase much more (0.8 ◦ C per decade) than those in equatorial regions (0.1 ◦ C per
decade). Global mean sea levels are expected to rise ca. 6 cm per decade, thus causing
immediate threats (increased salt water intrusion, loss of land and natural resources)
to coastal regions.
A rise of global temperatures has been predicted to be accompanied by increased
frequency and destructiveness of hurricanes, more protracted droughts, longer and
hotter heat waves, more severe rainy periods and significant changes in the area of
the great ice sheets of Antarctica (McNeely et al. 1995). However, there appears to
be no hard evidence to substantiate all these assertions (Mahlman 1997).

Biodiversity

The word ‘biodiversity’ is a contraction of ‘biological diversity’. Biodiversity has

been defined by many scientists and politicians associated with governmental and
non-governmental organizations (e.g. McNeely et al. 1990; Groombridge 1992;
Wilson 1992; Boyle and Boontawee 1995; Heywood and Watson 1995; Lammerts van
Bueren and Duivenvoorden 1996). In their pioneer policy document entitled ‘Con-
serving the World’s Biological Diversity’ McNeely et al. (1990) consider biological
diversity as a concept encompassing all species, plants, animals and micro-organisms
and the ecosystems and ecological processes of which they are parts. As they point
out, it is an umbrella term for the degree of nature’s variety, including both the number
and frequency of ecosystems, species or genes in a given assemblage. The 1992 Con-
vention on Biological Diversity defines biodiversity as ‘the variability among living
organisms from all sources, including, inter alia, terrestrial, marine and other aquatic
ecosystems and the ecological complexes of which they are part. This includes di-
versity within species (genetic diversity), between species (species diversity) and of
ecosystems (Heywood and Watson 1995).
A recent estimate of the number of species worldwide is 13 million, though only
about 1.6 million have actually been described (Heywood and Watson 1995). Certain
areas on the globe show exceptional concentrations of species with high levels of
endemism and unusually rapid rates of depletion (Myers 1988). These areas are called
‘hot spots’ and are found in certain tropical forests, coral reefs and Mediterranean
ecosystems (Myers 1988).
Major threats to biodiversity include habitat alteration and loss, over-harvesting,
chemical pollution, invasive species and increasing population pressure. Climate
change may modify and enhance local anthropogenic disturbances. According to
Jenkins (1992), rates of habitat modification are currently so high that virtually all nat-
ural terrestrial habitats and protected areas are destined to become ecological ‘islands’
in surrounding ‘oceans’ of habitat much altered. This process of fragmentation and
 1387

isolation – main concepts in island biogeographic theory (MacArthur and Wilson

1967) – is predicted to lead directly and indirectly to accelerated species extinctions at
both the local and the global scales, thus reducing the world’s biodiversity at all levels
(Jenkins 1992; Lawton and May 1995). Some authors (Myers 1979, 1989; Ehrlich
and Ehrlich 1981) argue that the loss of biodiversity threatens ecosystem integrity,
and may ultimately threaten human existence itself.

Effects of paleoclimate change on biodiversity

Understanding the impacts of past climate change on plant and animal life may fa-
cilitate the development of models that predict future shifts in species, communities
and ecosystems (Webb 1992; Lundqvist 1996; Lowe and Walker 1997). Over the
past million years, the most noticeable pattern of global climate variation has been
the oscillations between cold and warmer periods (glacials and interglacials), with
glacials occurring roughly every 100,000 years. According to Webb (1992) the past
18,000 years BP (Before Present), including the end of the last glacial, are particularly
interesting. The rise in global mean temperature of 5 ± 1 ◦ C during this period closely
approximates the 4.2 ± 1.2 ◦ C rise that is predicted for the near future, as a result of
a doubling of the effective concentration of greenhouse gases (Schlesinger 1989).
However, the rate at which the global mean temperature is predicted to rise under
future global warming conditions is faster than any natural warming during the past
18,000 years. This implies that many species are likely to be unable to move their
ranges rapidly enough to keep up with the changing climate (Webb 1992).
Studies in the Colombian Andes have shown the importance of Pleistocene
(2,000,000–10,000 years BP) climate change for environmental and floristic dynam-
ics along elevational gradients in tropical mountain regions (Van der Hammen 1989,
1992; Hooghiemstra and Cleef, 1995). Results depict strong altitudinal shifts in ve-
getation belts as a response to temperature changes. Palynological research in Costa
Rica has revealed a cooling of 7 to 8 ◦ C during the last glacial maximum (Islebe and
Hooghiemstra 1997). However, environmental change throughout Central America
during the mid Holocene (10,000–0 years BP) seems more affected by changes in
humidity than by temperature changes. Dendrochronological studies in Chile and the
northern hemisphere show the sensitivity of trees to temperature changes over the last
few hundred years (Peters 1992; Szeicz 1997).
Data from Africa stress the impacts of cooler and drier environments on paleotrop-
ical rain forest during the last ice age, about 70,000 to 12,000 years BP (Bonnefille
et al. 1990; Maley 1996). Fossil pollen records from East African mountains show a
descent of vegetation zones indicating a 6 ◦ C change in temperature (Van Zinderen
Bakker and Coetzee 1972). As Sosef (1994) points out, the area of lowland rain forest
presumably shrank considerably during the last glacial period and ultimately disin-
tegrated into a number of small refuges, situated as islands within an area occupied
by more drought resistant vegetation. African shade-loving rain forest species such
1388

as those belonging to the genus Begonia are presently concentrated within the main
postulated refuge localities. They can be regarded as dependable bio-indicators of
former refuges (Sosef 1994).

Effects of recent climate change on biodiversity

In this section a number of observations of effects of climate change on biodiversity

are described, i.e. (i) shifts of major vegetation zones or biomes, (ii) shifts in ranges
of individual species and in the composition of species assemblages, (iii) interactions
between effects of climate change and habitat fragmentation and (iv) changes in
ecosystem functioning.

(i) Shifts of major vegetation zones or biomes

Changes in global vegetation cover and in the boundaries of the world’s biomes are
expected to occur in response to global climate change (McNeely et al. 1990; Peters
and Lovejoy 1992; Heywood and Watson 1995). As the earth warms, species are gen-
erally expected to shift to higher latitudes (poles) and altitudes (peaks). For example,
the timber line in Finland would rise about 200 m higher into areas where it used
to be during the Atlantic thermal period about 4,500 to 7,500 years ago (Kellomäki
1996). Under projected future global warming conditions, certain areas of endemic
and species-rich tropic alpine vegetation may be fully replaced by montane cloud
forests presently found at lower altitudes (Halpin 1994).
However, models simulating the displacement of vegetation types or biomes have
been criticized for being too simplistic (Leemans 1996; Cramer 1997; Halpin 1997).
First, simulated patterns of shifts of biomes as complete entities are not very realistic,
because they do not take into account the individual responses of species to changes in
climate factors (Leemans 1996). In more recent models, e.g., BIOME (Prentice et al.
1992) plant types show an individualistic response to climate, and new assemblages
of plant types can develop in response to climate change. Second, the outcomes of
models are often presented as potential vegetation maps based on future equilibrium
climates, without an indication of the time required to reach this new stage (Halpin
1997). Third, predictions of poleward and altitudinal movements of species under
warmer climates are often based on temperature responses only. When moisture vari-
ables are taken into account, the results are far more complex. For example, Halpin
(1997) showed that changes in ecoclimatic zonation of mountain areas are generally
nonsymmetrical in response, and widely different between latitudes.

(ii) Shifts in ranges of individual species and in the composition of species

assemblages
Changes in climate may affect the physiology, phenology and interspecific interac-
tions between individual species, and as a consequence, shifts in geographic distri-
butions may occur. For example, range changes observed for several butterflies in
 1389

Britain are ascribed to small temperature increases (less than one degree) during this
century (Ford 1982). Similarly, the northward expansion of birch into the Swedish
tundra is attributed to warming during the first half of this century (Kullman 1983).
However, if affected species are not able to adjust their geographic distribution, their
survival chances will be strongly reduced.
In the Netherlands, over the last few decades, 64 vascular plant species have be-
come extinct, while about 84 species have appeared for the first time (Van der Meijden
1993). For many of these 84 ‘neophytes’, the northernmost limit of their geographic
range is now in the Dutch delta. Both climate change and direct human influences
on ecosystems are assumed to have caused these shifts in species distributions. A
study by Nabuurs et al. (1997) reports on the possible effects of climate change on
forest ecosystems. Current climate scenarios predict drastic changes in tree species
composition in the long term.
Recent shifts in phenology and distribution of a large sample of Microlepidoptera
in the Netherlands are also related to climate change (Ellis et al. 1997a, 1997b).
During the period from 1975 to 1994, the flight peak shifted on an average to a date
11.6 days earlier, presumably due to the rise in spring temperatures. More than 50%
of the species examined had undergone a significant change in distribution over the
same period.
Species that are especially sensitive to climate change may be used as indicator
species (‘bio-indicators’) for assessing the climate sensitivity of whole ecosystems.
De Groot et al. (1995) describe preliminary results on the selection of bio-indicators,
based on six criteria: climate sensitivity, habitat constraints, position within distribu-
tion range, dispersal capacity, functional position in the ecosystem and suitability for
monitoring. A number of herbaceous plant species, butterflies and birds are identified
as suitable bio-indicators for climate change in the Netherlands and Western Europe.
Shifts in geographic distributions of individual species and in the composition of
species assemblages can be identified by long-term monitoring studies, using, e.g.,
Geographic Information Systems (GIS) (Heil and Van Deursen 1996). Natural eco-
climatic transitions or ecotones may be especially suitable for monitoring effects of
climate change, because they are likely to be especially sensitive to climate change.
Examples are tropical tree border lines such as those found at the rainforest–savanna
boundary (De Wilde and Van der Maessen 1997) and cloud forest – paramo inter-
face (Cleef 1981; Kappelle et al. 1995) and temperate timber lines such as found in
northern Finland (Kellomäki 1996).
Examples of monitoring studies in marine environments are the studies on tropical
coral reefs by Bak and Nieuwland (1995), and that on sublittoral communities in the
North Sea by De Kluijver (1997). Bak and Nieuwland (1995) monitored permanent
quadrates for over two decades and showed a significant decrease in coral colonies,
particularly at disturbed shallower reefs. Whereas most of the degradation processes
are directly related to human influence, a rise in the temperature of ocean waters will
lead to drastic reef degradation in the long run. De Kluijver (1997) characterizes the
1390

structure, composition and synecology of a series of sublittoral communities. These

communities are expected to change as a consequence of climate change, resulting
in a sea level rise, warming of seawater, shifts in seasonal fluctuations and a rise in
carbon concentrations.

(iii) Interactions between effects of climate change and habitat fragmentation

Over the past decades, destruction and fragmentation of habitats has led to bio-
diversity depletion on local, regional and global scales (Harris 1984; Myers 1989).
Habitat fragmentation in conjunction with climate change sets the stage for an even
larger wave of extinction than previously imagined, based on consideration of human
encroachment alone (Myers 1989; Peters and Lovejoy 1992). Habitat fragmentation
results in isolation of species populations and may locally lead to a reduction of
genetic variation. Fragmentation may also prevent species migration and dispersal
towards more suitable habitats in response to climate change (MacArthur and Wilson
1967; Peters and Lovejoy 1992; Bierregaard et al. 1997).
In addition to its effects on migration of populations, fragmentation of landscapes
into smaller units can have a large impact on the climate along the edges and inside the
remaining fragments. In many forested areas, fragmentation has produced landscapes
in which forest edges are a dominant feature and edge influences are extensive in the
remaining tracts of the forest (Chen et al. 1993). Saunders et al. (1993) show that rain
forest remnants surrounded by cleared agricultural lands are exposed to markedly
different solar radiation, wind, water and nutrient regimes than continuously forested
regions. Kapos (1989) and Turton and Freiburger (1997) also measure changes in cli-
mate along forest interior-edge-exterior gradients in tropical rain forest. Biodiversity
may change significantly along these gradients, as is shown for vascular plants along
belt transects perpendicular to the edge of cloud forest fragments (Williams-Linera
1990; Oosterhoorn and Kappelle 1999).
In habitat fragments, survival of populations or species that need to adjust their
geographic distribution to a changing climate will depend on their migration potential,
but also on fragment size, quality and distribution (between-fragment distance, con-
nectivity via corridors), and the character of the matrix surrounding fragments (phys-
ical barrier type). Establishing corridors between fragments is an often-suggested
management response to changing climate conditions, though much is still unknown
about their potential effectiveness. Knowledge is lacking on the required spatial con-
figuration of corridors, e.g., their optimal width, edge to area ratios and orientation in
different environments (Halpin 1997).

(iv) Changes in ecosystem functioning

The biogeochemical functioning of an ecosystem depends on the summed, inter-
related activities of its organisms, i.e. the ways and rates at which they carry out
ecosystem processes (e.g., respiration, carbon dioxide fixation, nitrification, litter de-
composition). Climate change may influence ecosystem functioning if the physiology
 1391

of species is affected, e.g., by changes in temperature or moisture availability. Changes

in climate factors may also exceed the physiological tolerance of species and/or dis-
turb their functional relationships with others, causing species to become extinct or
migrate to other sites, and probably reducing the ecosystem’s biodiversity.
Current losses in biodiversity by human-induced environmental changes has re-
newed interest in research on the significance of biodiversity for ecosystem function-
ing and resilience with respect to stress and disturbance (Naeem et al. 1994; De Ruiter
et al. 1995; Folke et al. 1996; Mooney et al. 1996; Brussaard et al. 1997; Chapin
et al. 1997; Díaz and Cabido 1997; Hooper and Vitousek 1997; Tilman et al. 1996,
1997; Wardle et al. 1997). In his commentary on recent outcomes of field experiments
(Hooper and Vitousek 1997; Tilman et al. 1997; Wardle et al. 1997), Grime (1997)
concludes that there is currently no convincing evidence that ecosystem processes are
crucially dependent on higher biodiversity. Rather, the functional characteristics of
the dominant plant species would be important in controlling ecosystem processes.
However, it remains obvious that losses of species or functional groups from an
ecosystem could at some point impair its functioning and its capacity to provide ser-
vices to society (Mooney et al. 1996; Chapin et al. 1997; Grime 1997). Some species
or functional groups carry out unique functions and cannot be substituted by others.
But also the loss of species with similar ecosystem effects may reduce ecosystem
resilience and narrow the options for adjustments to climate change.

Identification of key research issues

(a) Interviews with research groups in the Netherlands

During interviews with ca. 30 biological research groups in the Netherlands, re-
search suggestions were made for conducting studies on effects of climate change
on biodiversity. These suggestions can be grouped into the following six key research
issues:

(i) Spatial and temporal distributions of taxa. This type of research seeks correl-
ations between long-term data on distributions of taxa and long-term climate data
within an area. By reconstructing past changes, potential future changes in the distri-
butions of taxa under climate change scenarios may be understood. Different sources
of information (field records, specimens from systematic collections, fossils) may be
used, depending on the time scale at which the research is undertaken.

(ii) Migration and dispersal potentials of taxa. Autecological research on a number

of plant and/or animal species (possibly representing functional groups of species)
may answer the following questions: How fast can species migrate in response to cli-
mate change? What are their chances for survival and establishment in new habitats?
and, what are the consequences of species migration for biodiversity in the abandoned
and the newly colonized habitats?
1392

(iii) Genetic diversity and viability of (meta)populations of species (including

research on the effects of habitat fragmentation). Climate change may result in
isolation of populations within a species, putting constraints on the maintenance
of genetic diversity, with the ultimate risk of extinction. Measurements of the ge-
netic diversity within a selected metapopulation will enable an assessment of the
degree of reproductive isolation and the viability of local populations, as well as the
consequences for the genetic diversity of the metapopulation in the long term.

(iv) Physiological tolerance of species. Changes in temperature or moisture avail-

ability will affect physiological processes in plants and animals. In addition, the
phenology of many organisms, i.e. the timing of different phases in their life cycles,
will be affected (e.g. the onset and duration of winter rest). Changes in climate factors
and their seasonal patterns, and the magnitude and timing of extreme values are im-
portant. Assessment of species’ tolerance for changes in climate factors will enable
assessment of the risks posed by climate change for the survival of those species. This
is especially relevant for (rare) species that do not have a high dispersal or migratory
capacity.

(v) Disturbance of functional interactions between species. Species are affected in

different ways by climate change, implying that functional interactions between spe-
cies will be affected. The consequences may be positive or negative for the survival
of individuals of either or both species (and possibly others). Numerous examples of
species interactions can be given, e.g. between plants and herbivores, between pred-
ators and preys, etc. Special cases of disturbances of functional interactions between
species involve ‘displacements’ in time and/or space, which are often caused by
phenological changes.

(vi) Ecosystem processes (including research on the direct effects of increased

concentrations of greenhouse gases such as carbon dioxide). Climate change may
affect the physiology, phenology and interspecific interactions of individual species
and the species composition of ecosystems. All these responses can have con-
sequences for ecosystem processes, e.g., primary production, food web relations, and
nutrient cycling. In turn, changes in ecosystem processes have complex feedback
effects on the functioning of species, species composition and possibly climate (e.g.,
increased production of carbon dioxide after warming and drying of tundra ecosys-
tems). Assessments of long-term consequences of changes in ecosystem processes
will require experimental work as well as the development of simulation models.

(b) Recommendations for an NRP research programme

The results of the interviews were used to formulate recommendations for a 2-year
research programme to be carried out within the Dutch National Research Programme
on Global Air Pollution and Climate Change (NRP). It was recommended to focus
the NRP programme on two questions, i.e.:
 1393

(1) What are the effects of current and potential climate change on spatial and
temporal distribution patterns and on the existence of species and ecosystems?

(2) What biological mechanisms are involved in the responses of species and eco-
systems to global climate change?
In addition, two complementary approaches were recommended, i.e.:
(i) Analyses of long-term data sets comprising biogeographical and climate obser-
vations and measurements;
(ii) Experimental work on a number of species or functional groups of species aimed
at investigating their physiological or phenological responses to climate change,
and their migration or dispersal potential.
The emphasis of the NRP programme was put on research in the Netherlands and
north western Europe, although it was realized that a major effort is needed to mon-
itor and predict the effects of climate change in the tropics. Research priorities for
the tropics will depend on the state of knowledge of genetic, taxic and ecosystem
diversity for a given region.

Concluding remarks

Assessing the impact of climate change on biodiversity is difficult, due to the spatial
and temporal scale and the complexity of the problem, and its interactions with other
environmental factors. This is illustrated by the broad range of research issues iden-
tified in this paper. Recommendations for the research programme mentioned above
were based on these issues, but also directed by budgetary and time constraints of the
specific programme. Hence, another approach may be taken under different circum-
stances. However, a research programme should be interdisciplinary, and generate
input data for predictive models based on climate change scenarios.

Acknowledgements

This project was carried out in the framework of the Dutch National Research Pro-
gramme on Global Air Pollution and Climate Change; registered under no. 952250,
entitled: ‘Programming study: Biodiversity and Global Climate Change’. We thank
Jan van Groenendael, Wouter Los, Marco Roos, Wilko Verweij and the interviewed
scientists for their contributions.

References

Bak RPM and Nieuwland G (1995) Long-term change in coral communities along depth gradients over
leeward reefs in The Netherlands Antilles. Bulletin of Marine Science 56: 609–619
1394

Bierregaard RO Jr, Laurance WF, Sites JW Jr, Lynam AJ, Didham RK, Andersen M, Gascon C, Tocher
MD, Smith AP, Viana VM, Lovejoy TE, Sieving KE, Kramer EA, Restrepo C and Moritz C (1997)
Key priorities for the study of fragmented tropical ecosystems. In: Laurance WF and Bierregaard RO Jr,
(eds) Tropical Forest Remnants: Ecology, Management, and Conservation of Fragmented Communities,
pp 515–525. University of Chicago Press, Chicago, IL
Bonnefille R, Roeland JC and Guinot J (1990) Temperature and rainfall estimates for the past 40,000 years
in equatorial Africa. Nature 346: 347–349
Boyle TJB and Boontawee B (1995) Measuring and Monitoring Biodiversity in Tropical and Temperate
Forests. CIFOR Bogor, Indonesia
Brussaard L, Behan-Pelletier VM, Bignell DE, Brown VK, Didden W, Folgarait P, Fragoso C, Wall-
Freckman D, Gupta VVSR, Hattori T, Hawksworth DL, Klopatek C, Lavelle P, Malloch DW, Rusek
J, Söderström B, Tiedje JM and Virginia RA (1997) Biodiversity and ecosystem functioning in soil.
Ambio 26: 563–570
Bush M (1997) Ecology of a Changing Planet. Prentice Hall, Upper Saddle River, NJ
Chapin FS III, Walker BH, Hobbs RJ, Hooper DU, Lawton JH, Sala OE and Tilman D (1997) Biotic
control over the functioning of ecosystems. Science 277: 500–504
Chen J, Franklin JF and Spies TA (1993) Contrasting microclimates along clearcut edge and interior of
old-growth Douglas-fir forest. Agricultural and Forest Meteorology 63: 219–237
Cleef AM (1981) The Vegetation of the Páramo of the Colombian Cordillera Oriental. Dissertationes
Botanicae 61. Cramer, Vaduz, LI
Cramer W (1997) Using plant functional types in a global vegetation model. In: Smith TM, Shugart HH
and Woodward FI (eds) Plant Functional Types: Their Relevance to Ecosystem Properties and Global
Change, pp 271–288. IGBP. Cambride University Press, Cambridge, UK
Dale VH (1997) The relationship between land-use change and climate change. Ecological Applications
7: 753–769
De Groot RS, Ketner P and Ovaa AH (1995) Selection and use of bio-indicators to assess the possible
effects of climate change in Europe. Journal of Biogeography 22: 935–943
De Kluijver MJ (1997) Sublittoral communties of North Sea hard-substrata. PhD Thesis, University of
Amsterdam, Amsterdam
De Ruiter PC, Neutel A-M and Moore JC (1995) Energetics, patterns of interaction strengths, and stability
in real ecosystems. Science 269: 1257–1260
De Wilde JJFE and Van der Maessen LJG (1997) Diversity and Distribution of Caesalpinioid Species in
the Lopé Reserve in Gabon: A Search for Patterns and Their Explanation. NWO Proposal. Wageningen
Agricultural University, Wageningen, The Netherlands
Díaz S and Cabido M (1997) Plant functional types and ecosystem function in relation to global change.
Journal of Vegetation Science 8: 463–474
Ehrlich P and Ehrlich A (1981) Extinction: The Causes and Consequences of the Disappearance of Species.
Random House, New York, NY
Ellis WN, Donner JH and Kuchlein JH (1997a) Recent shifts in phenology of Microlepidoptera, related to
climatic change (Lepidoptera). Entomologische Berichten Amsterdam 57: 66–72
Ellis WN, Donner JH and Kuchlein JH (1997b) Recent shifts in distribution of Microlepidoptera in The
Netherlands. Entomologische Berichten Amsterdam 57: 119–125
Folke C, Holling CS and Perrings C (1996) Biological diversity, ecosystems, and the human scale.
Ecological Applications 6: 1018–1024
Ford M (1982) The Changing Climate. Georg Allen & Unwin, London
Grime JP (1997) Biodiversity and ecosystem function: the debate deepens. Science 277: 1260–1261
Groombridge B (ed) (1992) Global Biodiversity: Status of the Earth’s Living Resources. Compiled by the
World Conservation Monitoring Centre. Chapman and Hall, London
Halpin PN (1994) Latitudinal variation in the potential response of mountain ecosystems to climate change.
In: Beniston M (ed) Mountain Environments in Changing Climates, pp 180–203. Routledge, London
Halpin PN (1997) Global climate change and natural-area protection: management responses and research
directions. Ecological Applications 7: 828–843
Harris LD (1984) The Fragmented Forest. The University of Chicago Press, Chicago, IL
 1395

Heil GW and Van Deursen WPA (1996) Searching for patterns and processes: modelling of vegetation
dynamics with geographical information systems and remote sensing. Acta Botanica Neerlandica 45:
543–556
Heywood VH and Watson RT (eds) (1995) Global Biodiversity Assessment. UNEP. Cambridge University
Press, Cambridge, UK
Hooghiemstra H and Cleef AM (1995) Pleistocene climate change and environmental and generic dy-
namics in the Northern Andean montane forest and páramo. In: Churchill SP, Balslev H, Forero E and
Luteyn JL (eds) Biodiversity and Conservation of Neotropical Montane Forests, pp 35–49. The New
York Botanical Garden, Bronx, NY
Hooper DU and Vitousek PM (1997) The effects of plant composition and diversity on ecosystem
processes. Science 277: 1302–1305
Houghton JT (ed) (1995) The Science of Climate Change. Cambridge University Press, Cambridge, UK
Houghton JT, Jenkins GJ and Ephraums JJ (1990) Climate Change: The IPCC Scientific Assessment.
Cambridge University Press, Cambridge, UK
Houghton JT, Callander BA and Varney SK (eds) (1992) Climate Change 1992: The Supplementary Report
to the IPCC Scientific Assessment. Cambridge University Press, Cambridge, UK
Houghton JT, Meira Filho LG, Callander BA, Harris N, Kattenberg A and Mashell K (eds) (1996) Climate
Change 1995: The Science of Climate Change. Cambridge University Press, Cambridge, UK
Islebe GA and Hooghiemstra H (1997) A contribution to Late Quaternary vegetation and climate history
of Costa Rica. Quaternary Science Reviews 16: 589–604
Jenkins M (1992) Species Extinction. In: Groombridge B (ed) Global Biodiversity: Status of the Earth’s
Living Resources, pp 192–205. Compiled by the World Conservation Monitoring Centre (WCMC),
Chapman and Hall, London
Kapos V (1989) Effects of isolation on the water status of forest patches in the Brazilian Amazon. Journal
of Tropical Ecology 5: 173–185
Kappelle M, Kennis PAF and De Vries RAJ (1995) Changes in diversity along a successional gradient in
a Costa Rican upper montane Quercus forest. Biodiversity and Conservation 4: 10–34
Kellomäki S (1996) Effects of changes in climate, forest use and forestry practices on forest structure
and the biodiversity of boreal forests. In: Bachmann P, Kuusela K and Uuttera J (eds) Assessment of
Biodiversity for Improved Forest Management, pp 53–58. European Forest Institute Proceedings No. 6,
European Commission DG XII, Brussels
Kullman L (1983) Past and present tree lines of different species in the Handolan Valley, Central
Sweden. In: Morisset P and Payette S (eds) Tree Line Ecology, pp 25–42 Centre d’Etude Nordiques
de l’Université Laval, Québec
Lammerts van Bueren EM and Duivenvoorden JF (1996) Towards Prioirities of Biodiversity Research in
Support of Policy and Management of Tropical Rain Forests. The Tropenbos Foundation, Wageningen,
The Netherlands
Lawton JH and May RM (eds) (1995) Extinction Rates. Oxford University Press, Oxford
Leemans R (1996) Biodiversity and global change. In: Gaston KJ (ed) Biodiversity: A Biology of Numbers
and Difference, pp 367–387. Blackwell Science, Oxford
Lowe JJ and Walker MJC (1997) Reconstructing Quaternary Environments. 2nd ed. Longman Group Ltd,
Essex, UK
Lundqvist J (1996) Quaternary climatic fluctuations, global environment changes and the impact of man.
Nature and Resources 32: 30–37
MacArthur RH and Wilson EO (1967) The Theory of Island Biogeography. Princeton University Press,
Princeton, NJ
Mahlman JD (1997) Uncertainities in projections of human-caused climate warming. Science 278: 1416–
1417
Maley J (1996) The African rain forest – main characteristics of changes in vegetation and climate from
the Upper Cretaceous to the Quaternary. Proceedings of the Royal Society of Edinburgh 104 B: 31–73
McNeely JA, Miller KR, Reid WV, Mittermeier RA and Werner TB (1990) Conserving the World’s Bio-
logical Diversity. IUCN, Gland, Switzerland; WRI, CI, WWF-US, Washington, DC, The World Bank,
Washington, DC
1396

McNeely JA, Gadgil M, Leveque C, Padoch C and Redford K (eds) (1995) Human influences on
biodiversity. In: Heywood VH and Watson RT (eds) Global Biodiversity Assessment, pp 711–821.
Cambridge University Press, UNEP. Cambridge, UK
Mooney HA, Cushman JH, Medina E, Sala OE and Schulze E-D (eds) (1996) Functional Roles of
Biodiversity: A Global Perspective. ICSU-UNEP. John Wiley, Chichester, UK
Myers N (1979) The Sinking Arc. Pergamon Press, Oxford
Myers N (1988) Threatened biotas: ‘Hot spots’ in tropical forests. The Environmentalist 8: 187–208
Myers N (1989) Deforestation Rates in Tropical Forests and Their Climate Implications. Friends of the
Earth, London
Nabuurs GJ, Kramer K and Mohren GMJ (1997) Effecten van Klimaatverandering op het Neder-
landse Bos en Bosbeheer. Report 256. DLO Institute for Forestry and Nature Research, Wageningen,
The Netherlands
Naeem S, Thompson LJ, Lawler SP, Lawton JH and Woodfin RM (1994) Declining biodiversity can alter
the performance of ecosystems. Nature 368: 734–737
Oosterhoorn M and Kappelle M (1999) Vegetation structure and composition along an interior-edge-
exterior gradient in a Costa Rican montane cloud forest. Forest Ecology and Management (in
press)
Peters R (1992) Ecology of Beech Forests in the Northern Hemisphere. PhD Thesis, Wageningen
Agricultural University, Wageningen, The Netherlands
Peters RL and Lovejoy TE (1992) Global Warming and Biological Diversity. Yale University Press,
New Haven, CT, 386 pp
Prentice IC, Cramer W, Harrison SP, Leemans R, Monserud RA and Solomon AM (1992) A global biome
model based on plant physiology and dominance, soil properties and climate. Journal of Biogeography
19: 117–134
Saunders DA, Hobs RJ and Arnold GW (1993) The Kellerberrin project on fragmented landscapes: A
review. Biological Conservation 5: 18–32
Schlesinger ME (1989) Model projections of the climate changes induced by increased atmospheric CO2 .
In: Duplessy JC, Berger A and Schneider SH (eds) Climate and Geosciences, pp 375–415. Kluwer
Academic Publishers, Dordrecht, The Netherlands
Sosef MSM (1994) Studies in Begoniaceae. V. WAU Papers 94.1. Wageningen Agricultural University,
Wageningen, The Netherlands
Sprengers SA, Slager LK and Aiking H (1994) Biodiversity and Climate Change. Part I: Establishment
of Ecological Goals for the Climate Convention. Free University Institute for Environmental Studies.
Amsterdam
Szeicz JM (1997) Growth trends and climate sensitivity of trees in the North Patagonian rain forest of
Chile. Canadian Journal of Forestry Research 27: 1003–1014
Tilman D, Wedin D and Knops J (1996) Productivity and sustainability influenced by biodiversity in
grassland ecosystems. Nature 379: 718–720
Tilman D, Knops J, Wedin D, Reich P, Ritchie M and Siemann E (1997) The influence of functional
diversity and composition on ecosystem processes. Science 277: 1300–1302
Turton SM and Freiburger HJ (1997) Edge and aspect effects on the microclimate of a small tropical
forest remnant on the Atherton Tableland, Northeastern Australia. In: Laurance WF and Bierregaard RO
(eds) Tropical Forest Remnants: Ecology, Management, and Conservation of Fragmented Communities,
pp 45–54. University of Chicago Press, Chicago, IL
Van der Hammen T (1989) History of the montane forests of the northern Andes. Plant Systematics and
Evolution 162: 109–114
Van der Hammen T (1992) Global change, shifting ranges and biodiversity in plant ecosystems. In: Sol-
brig OT, Van Emden HM and Oordt PGWJ (eds) Biodiversity and Global Change, pp 159–166. IUBS
Monograph 8. IUBS, Paris
Van der Meijden R (1993) Proefproject Flora en Fauna 2030. Series ‘Achtergrondreeks’, Vol 2. FLORON
Foundation and Ministry of Public Housing Spatial Planning and Environment (VROM), The Hague
Van Vuuren MMI and Kappelle M (1998) Biodiversity and Global Climate Change. Report no.
410.200.014, Dutch National Research Programme on Global Air Pollution and Climate Change.
National Institute for Public Health and the Environment (RIVM), Bilthoven, The Netherlands
 1397

Van Zinderen Bakker EM and Coetzee JA (1972) A re-appraisal of Late Quaternary climate evidence from
tropical Africa. In: van Zinderen Bakker EM (ed) Palaeoecology of Africa, Vol 7, pp 151–191. A.A.
Balkema, Rotterdam, The Netherlands
Wardle DA, Zackrisson O, Hörnberg G and Gallet C (1997) The influence of island area on ecosystem
properties. Science 277: 1296–1299
Watson RT, Zinyowera MC, Moss RH and Dokken DJ (eds) (1996) Climate Change 1995, Impacts,
Adaptations and Mitigation of Climate Change: Scientific–Technical Analyses. IPCC. Cambridge
University Press, Cambridge, UK
Webb T (1992) Past changes in vegetation and climate: Lessons for the future. In: Peters RL and Lovejoy
TE (eds) Global Warming and Biological Diversity, pp 59–75. Yale University Press, New Haven, CT
Williams-Linera G (1990) Vegetation structure and environmental conditions of forest edges in Panama.
Journal of Ecology 78: 356–373
Wilson EO (1992) The Diversity of Life. Allen Lane – Penguin, London
Zwerver S, Van Rompaey RSAR, Kok MTJ and Berk MM (1995) Climate Change Research: Evaluation
and Policy Implications. Studies in Environmental Science 65 (B). Elsevier, Amsterdam