POLICY PERSPECTIVE
Ecosystem management in Madagascar during global change
Malika Virah-Sawmy
Oxford University Centre for the Environment, University of Oxford, South Parks Road, Oxford OX 1 3QY, U.K.
Keywords
Biodiversity offset; climate change; drought;
ecotone; mining; mosaic; refuge.
Correspondence
Malika Virah-Sawmy, WWF Madagascar and
Western Indian Ocean Programme Office,
B.P. 738, (101) Antananarivo, Madagascar.
Tel: +261 20 22 348 85; fax: +261 20 22 348 88.
E-mail: [email protected]
Received: 14 November 2008; accepted 3 May
2009.
doi: 10.1111/j.1755-263X.2009.00066.x
Abstract
A long-held assumption about Madagascar is that all its open vegetation and
forest fragments represent anthropogenically degraded landscapes. Further, its
forests are believed to lack regenerative capacity following extirpation. This
article presents a different ecological framework for southeast Madagascar, a
region with diverse forest habitats (humid, littoral, and transitional forests)
and exceptional biodiversity. Recently published reconstructions of vegetation
change at millennial scales indicate that the mosaic vegetation in southeastern Madagascar represents a natural distribution, with littoral forest spatially
constrained by soil moisture, rather than principally fragmented by human
activity. There is also evidence that the littoral forest is resilient to climatic
change and has acted as refuges and centers of diversification for millennia.
By contrast, Uapaca woodlands, which connected the littoral forest fragments,
switched to a stable fire-maintained Ericoid grassland in response to climatic
changes. Evidence also suggests that local communities managed forest resources sustainably in the past, but human migration, insecure land tenure,
and road expansion may have accelerated forest loss over the last decades. This
novel perspective demonstrates the integrated nature of ecological, geomorphological, climatical, and social systems. Finally, combining long-term with
contemporary perspectives, some strategies for maintaining biodiversity under
global change, including anthropogenic climate change and mining pressures
(and associated biodiversity offset projects) are suggested.
Introduction
Madagascar is one of the world’s top conservationpriority areas (Mittermeier et al. 2005) with estimates suggesting that its exceptional biota harbors around 3% of
global plant and vertebrate diversity (Myers et al. 2000).
Within the literature on Madagascar, there is a longheld assumption that 90% of Madagascar’s original forest cover has been destroyed by fires and slash-and-burn
agriculture (e.g., Hannah et al. 2008). Further, it has also
long been assumed that forests and woodlands covered
90% or more of presettlement Madagascar (Humbert &
Cours Darne 1965). Forest “remnants” and open vegetation (e.g., grasslands, savannas, and lowland Ericoid
grasslands) are therefore viewed as indicative of anthropogenically degraded. Both assumptions are based on the
apparent poverty of species and endemism in the open
vegetation relative to the forested parts of Madagascar
(Lowry et al. 1997). For example, it is estimated that more
than 90% of Madagascar’s endemic animal species live
exclusively in the forests and woodlands (Dufils 2003).
Moreover, there are indeed several accounts of deforestation since the French control of the island from 1890, although forest loss that has occurred cannot be attributed
directly to rapid population growth (see details in Jarosz
1993). Data on past forest cover prior to 1950 are speculative at best, but there is clearer evidence from the 1950s.
A recent study indicated that 27% of Madagascar was
forested around 1950 and forest cover has further declined to approximately 16% around 2000 (Harper et al.
2008); deforestation was particularly severe in the southwestern spiny forests, lowland evergreen forests, and
the western deciduous forests (Green & Sussman 1990;
Harper et al. 2008). Although the whole forest–island
c
Conservation Letters 2 (2009) 163–170 Copyright and Photocopying: 2009
Wiley Periodicals, Inc.
163
Ecosystem management in Madagascar
M. Virah-Sawmy
Figure 1 Distribution of littoral forest (area in hectares
in parentheses) along the eastern coast (bioclimatic
map modified from (Cornet 1974) and distributional
and littoral forest extent in 2005–2006 from Missouri
Botanical Garden and Vincelette et al. (2007)).
idea has been hotly disputed (Burney 1987; Lowry et al.
1997), it is surprising that little is known on the antiquity of mosaics in Madagascar; that is, whether they are
natural or human-modified systems, and processes maintaining these forest–grassland mosaics. A lack of empirical
evidence not only provides a weak basis for an ecological understanding of landscape patterns and processes in
Madagascar, most importantly, it can lead to misguided
conservation and land-use planning.
An understanding of ecosystem dynamics and spatial
patterns is especially critical in regions, which harbor substantial portions of the island’s biodiversity. The southeast
is one of the regions in Madagascar that support exceptional biodiversity and endemism as well as a high spatial turnover (beta diversity) unparalleled in almost any
part on the island (Goodman & Ramanamanjato 2007).
Within a short distance of the littoral forest (Figure 1)
on the southeast coast there are humid habitats spanning from montane to lowland forest descending on the
eastern mountains and transitional forest and dry spiny
164
bushlands towards the south (Figure 2A). Wooded grasslands currently separate these highly diverse forest habitats. But did grasslands always isolate these habitats? How
did climate changes shift the geographical ranges of these
disparate habitats? And, consequently, should we conserve them using a static or dynamic conservation framework?
It is hypothesized that prior to anthropogenic deforestation, the littoral forest connected a range of disparate
habitats by being contiguous with the lowland forest on
the eastern mountains and being continuous from north
to south along the eastern coast (de Gouvenain & Silander 2003; Consiglio et al. 2006). The littoral forest exists today as a series of species-rich humid to subhumid
fragments along the east coast (Figure 1). This ecosystem is widely seen as national priority for conservation
(Ganzhorn et al. 2001) due to its limited spatial extent,
currently estimated to cover 500 km2 (Du Puy & Moat
2003), high concentrations of range-restricted endemic
plants (at least 25% only occur in this habitat type)
Conservation Letters 2 (2009) 163–170
c
Copyright and Photocopying: 2009
Wiley Periodicals, Inc.
Ecosystem management in Madagascar
M. Virah-Sawmy
Figure 2 (A) Map of the southeastern Madagascar showing a diversity
of forest habitats (littoral, humid, transitional, spiny forests) interspersed
with a matrix of wooded or Ericoid grassland (in white) in relation to sedimentary sequences collected (at sites S9, W, M15, Z). (B) Summary findings
of long-term vegetation reconstruction in two littoral forest fragments and
their surrounding vegetation (at sites S9, W, M15, Z) at millennial scales
modified from Virah-Sawmy et al. (2009). Palaeoecological data indicate
that the littoral forest (sites S9,M15) was resilient to climate change (aridity
and marine inundations) in contrast to the surrounding Uapaca woodlands
(sites W, Z). (C) Long-term data suggest that the current vegetation mosaic
in the southeast is a natural distribution (Photographs: site S9 (depicting
littoral forest edge/Ericoid grasslands), site X (woody grasslands between
Mandena and Ste-Luce)), site M15 (depicting littoral forest interior).
(Consiglio et al. 2006), amphibians, reptiles, and birds,
and a unique assemblage of dry and humid taxa. The
large number of local endemics distributed within the littoral forest fragments (the smallest forest ecosystem on
the island) have led to speculations that the littoral forest
is a relict habitat and that 90% of the littoral forest has
been lost by human activities (Consiglio et al. 2006).
Whether the eastern coast of Madagascar was entirely
forested by littoral forests is not merely of interest for
environmental historians, it also has real implications in
the way we perceive and manage landscapes. For example, the view that may have influenced the clearance of
the southern littoral forest for mining is that this forest has been declining from prehistoric to present times
due to anthropogenic activities (Consiglio et al. 2006)
and that at current rates of deforestation this ecosystem
will be extinct by the year 2040 even in the absence of
mining (QIT Madagascar Minerals S.A. 2001). This article challenges the prevailing ecological and conservation
paradigm in Madagascar by reviewing recently published
long-term reconstructions of vegetation in the southeast
region. The new ecological framework for the southeast
presented here encompasses the influence of habitat mosaics, episodic climatic changes, differential resilience to
climatic changes, and recent deforestation and its complex and inter-linked drivers. The long-term perspective adds crucial information for biodiversity conservation
under global change—forthcoming climate change and
c
Conservation Letters 2 (2009) 163–170 Copyright and Photocopying: 2009
Wiley Periodicals, Inc.
165
Ecosystem management in Madagascar
M. Virah-Sawmy
current human impacts such as an opencast mining operation by QIT Madagascar Minerals (QMM), a subsidiary
of Rio Tinto. This perspective will contribute to Madagascar’s national vision—“Madagascar Naturally”—which
aims for “the development and implementation of environmental best practice” (MAP 2006).
Habitat mosaics
Occurrences of mosaics of forest–grassland in regions that
climatically can support forests are commonly attributed
to previous human activity (Willis et al. 2008). It has been
shown through numerous palaeoecological analyses that
this assumption is correct in America and northwest Europe (Willis et al. 2008). What is less well known, however, is the role that humans have played in the formation of such mosaics in tropical environments such as
Madagascar.
A set of recently published palaeoecological data from
multiple sites in the southeastern region represents the
first attempt to reconstruct mosaic dynamics in Madagascar. These vegetation reconstructions spanning the last
6,500 years in four habitats (two littoral forest fragments
and surrounding vegetation) have demonstrated that the
interface of littoral forest and its surrounding vegetation
composed currently of Ericoid grasslands predates human arrival on the island around 2,000 years ago (VirahSawmy et al. 2009, in press) (Figure 2). Although the
whole eastern portion of the island is part of one humid bioclimatic zone (Figure 1), these reconstructions
have shown that the distribution of littoral forest has
nevertheless been spatially constrained by soil moisture,
which is itself influenced by groundwater (Virah-Sawmy
et al. 2009, in press). The boundaries of littoral forest appear to have been relatively stable, adjoining temporally
shifting Uapaca woodlands—drought induced and fire
maintained Ericoid grasslands (see subsequently). These
results therefore contradict estimates of 90% littoral forest loss by Consiglio et al. (2006) because their studies
of the past distribution of littoral forest was based on
an extrapolation on extent of sandy soils, ignoring finescale soil moisture, which is probably the most important variable determining the occurrence of the littoral
forest. It is therefore suggested that positive feedback
mechanisms between abiotic (e.g., soil moisture, fires,
and drought) and biotic factors have created more than
one stable alternative system in southeastern Madagascar. There is indeed growing evidence for alternative stable states through time and space (e.g., Folke et al. 2004).
For example, it has been shown that rainforest/fireprone mosaics in the wet tropics of Australia—a mosaic
landscape comparable to some regions of Madagascar—
166
are a consequence of the interaction of local vegetation
positive feedback loops with the region’s environmental parameters (Warman & Moles 2009). Alternative stable states may explain the antiquity of the mosaic in
Madagascar.
Another key biogeographical question for Madagascar is the type of vegetation that connected the littoral
forests. Further, what was the vegetation that connected
the littoral fragments to the rainforest on the eastern
mountains (Anosyenne and Vohimenas ranges)? Paleoecological records from two sites in the surrounding
vegetation some distance away from the littoral forest
indicate that open to semi-open Uapaca/Myrica or Uapaca woodlands were once extensive there (Virah-Sawmy
et al. 2009, in press), possibly extending from the coast
to the lowland rainforests along the eastern coast. Interestingly, these woodlands were no-analog communities (communities that were compositionally unlike any
found today) containing a number of shrubs such as Canthium and Allophyllus and herbs such as several members of the Solanaceae family, grasses, and sedges. Currently, Uapaca-Sarcolaenaceae “Tapia” woodland exists
in Madagascar only in the southern central highlands
where it is assumed to be a relict vegetation that has been
transformed elsewhere to grasslands through regular
human-made fires (Du Puy & Moat 2003). In the “Tapia”
woodland, the dominant Uapaca species is U. bojeri, a firetolerant species. In contrast, palaeoecological evidence
from the southeast indicates that the Uapaca species of the
southeast woodlands were fire-intolerant, indicating an
affinity to Madagascar’s Uapaca forest clades. It is possible
that these woodlands were dominated by any of the five
Uapaca from the littoral forest including two littoral forest
endemics, U. ferruginea, U. littoralis, and three Madagascar
endemics, U. densifolia, U. louvelii, and U. sp. In sum, it is
plausible that Uapaca woodlands were once much more
extensive in Madagascar.
However, it appears that the Uapaca–Myrica woodland
was transformed in the region of Ste-Luce (southeast
Madagascar) around 5,800 calibrated years before present
(cal. yr BP) into Ericoid grasslands in response to climatic desiccation (Virah-Sawmy et al. 2009, in press)
(Figures 2 and 3). The Ericoid grasslands in Ste-Luce persisted to present time as naturally fire-maintained systems in the absence of people in the southeast for nearly
four millennia (Virah-Sawmy et al. 2009, in press). By
contrast, the open Uapaca woodlands of Mandena, further south of Ste-Luce, remained forested, acting possibly as a dispersal corridor with the humid forests on the
Vohimenas Mountains for several millennia until 1,000
cal. yr BP (Figures 2 and 3). It is suggested here that
the transformation of Uapaca woodlands into a more unsuitable matrix (i.e., for the dispersal of forest species)
Conservation Letters 2 (2009) 163–170
c
Copyright and Photocopying: 2009
Wiley Periodicals, Inc.
Ecosystem management in Madagascar
M. Virah-Sawmy
Figure 3 Schematic drawing showing Contraction and expansion of littoral forest and the retreat of dispersal corridors in response to different
environmental and anthropogenic factors over the last 6,500 years.
of Ericoid grasslands would have affected landscape
permeability, leaving populations of organisms with limited dispersal abilities (e.g., plants) as island populations
in some littoral forests, and more mobile ones (e.g., birds
and some lemurs) as part of a spatially extended population (patchy distribution) or metapopulation (regional
population persisting as the result of a balance between
the processes of local population extinction and patch
migration).
On a longer evolutionary time scale, it is suggested
here that natural contraction and expansion of Uapaca
woodlands, isolating and connecting the littoral forests to
each other and to the humid eastern rainforest for periodic intervals may explain the high floristic diversity
of the littoral forest. It also elucidates why these fragments are differentiated from each other. Given the recent formation of geological deposits on which the littoral forest rest dated between 150,000 to 80,000 years
BP (QIT Madagascar Minerals 2001), allopatric speciation through vicariant events during climatic desiccation
may explain why so many range-restricted local endemic
plants, amphibians, and reptiles evolved so quickly in
the littoral forest. These range-restricted species may also
have evolved prior to the formation of the littoral forest;
they may later on have become confined to these forests
due to past environmental changes.
Another interesting aspect of the littoral forest is that
it spans the transition from humid to transitional forest (Figure 1). The interface of littoral forest fragments
along a regional ecotone would have maximized local
and regional species richness and possibly facilitated diversification of a number of organisms giving rise to high
turnover in the southeast region (Goodman & Ramanamanjato 2007). There is mounting evidence that such areas of ecological discontinuities are adaptive components
for diversity, evolution, and persistence in variable environments (Noss 2001; Pressey et al. 2007).
Climate change response
The structure of an ecosystem often carries the signature
of episodic large-scale climatic changes and other disturbances, and it is important to take environmental history into account when interpreting present-day ecosystem dynamics. Further, long-term temporal perspectives
are essential for the development of meaningful conservation strategies that account for global change (Willis &
Birks 2006).
A recent study modeling species range shift in Madagascar to forthcoming climatic change predicted that the
littoral forest will disappear (Hannah et al. 2008). By
contrast, palaeoecological reconstructions show that the
littoral forest remained stable throughout several pronounced arid intervals, lasting hundreds of years each
during the last 6,500 years, as well as past sea-level rise of
1–3 m (Virah-Sawmy et al. 2009, in press), although temperature rises were not accounted for in this time-frame.
The littoral forests did, however, experience range contraction when sea level rise and associated marine surges
coincided with a severe drought 1,000 years ago. The
vegetation of the southeast coast shifted abruptly from a
mosaic vegetation (littoral forest, interspersed with both
Uapaca woodlands and Ericoid grasslands), into a uniform, open landscape of Ericoid grasslands with severely
reduced littoral forests (Figure 2B). There is also evidence from the fossil pollen records to indicate some plant
extinctions in this period (Virah-Sawmy et al. 2009, in
press).
The littoral forest, however, was more resilient and
recovered to near former levels within a few centuries,
although there was some community reassembly (e.g.,
there was an increase in generalist trees at the expense of
former dominants such as Symphonia, Dypsis, and Cynometra) (Virah-Sawmy et al. 2009, in press). The number of
range-restricted plant, amphibian, and reptile species that
c
Conservation Letters 2 (2009) 163–170 Copyright and Photocopying: 2009
Wiley Periodicals, Inc.
167
Ecosystem management in Madagascar
M. Virah-Sawmy
still exist in the isolated littoral forest indicate that it acted
as a refuge during climatic stress. The high resilience
of the littoral forest is attributed to forest–wetland systems retaining and reinstating former soil moisture levels
that allowed trees to recover and regenerate. By contrast,
the former Uapaca woodlands persisted as fire maintained
Ericoid grasslands possibly due to the drier soils. It
therefore appears that the adaptive capacity of the former open Uapaca woodlands is more easily exceeded than
littoral forest. Conservationists should therefore seek to
integrate features that can act as buffers against climate
change impacts, for example, forest - freshwater systems.
Two conclusions emerge: (1) landscape mosaic predates human arrival and this mosaic shows differences in
terms of resilience to climatic disturbances; (2) in conservation terms, the species-rich littoral forest shows greater
resilience and is reverting towards past species composition with high regenerative capacity. One of the most important implications emerging from these studies is that
organisms with limited dispersal abilities are less likely
to migrate from the littoral forest in response to forthcoming climate change, possibly due to the unsuitable
matrix around these forests. Nonetheless, the littoral forest has acted as refuges under stressful environmental conditions, and may possibly continue to do so if
they are not deforested; therefore, in-situ conservation is
important.
Socios-ecological systems
It is now recognized that indigenous people are neither
decoupled from, nor in complete balance with nature
(Malthusian vs. the Romanticism view) (Dove 2006). In
Madagascar, it is probable that many of the scattered littoral forest fragments provided scarce forest resources for
the Anosy on the eastern coast, and hence, may not have
been deforested. An alternative hypothesis is that the
low nutrient sandy soils on which the littoral forest rests
were not favored for agriculture in comparison to the humid forests on the lowland slopes (P.P. Lowry II, pers.
comm.). In general, local uses of the southern littoral forest (e.g., for medicinal plants, fuel wood, and construction) by the Anosy appear to have had less impacts on
the tree community over the last thousand years than
the impacts of climate change, although there were some
large forest fires at the peak of cultural transformation
during the Tranovato phase 400 years ago (Virah-Sawmy
et al. 2009, in press). Similarly, it has been demonstrated
that in contemporary times, local uses of the forests
have less impacts than the practices of migrant groups
(Ingram et al. 2005; Ingram & Dawson 2006) who may
have contributed to much of the deforestation observed
168
since the 1970s. Major human migration within Madagascar has been an important part of the island’s settlement history, and has often been induced by famine
and drought. For example in the extreme arid south, it
is well-documented that the droughts of 1970s, 1982,
1990–1992, and 2003 have resulted in declines in livestock numbers and grazing intensity causing large-scale
migration of people (Elmqvist et al. 2007). On one hand,
these droughts have allowed for more regeneration of
the tropical dry spiny forest given the declining pressures
(Elmqvist et al. 2007), but at the same time, may have
resulted in conflicts and depletion of natural resources in
regions where migrants took refuge.
Two important studies detailing deforestation rates in
the southern littoral forest using satellite imagery indicate that deforestation has been even more severe since
1992 (Ingram & Dawson 2006; Vincelette et al. 2007). An
analysis of the underlying causes of deforestation suggests
that deforestation by migrant groups may have also followed the construction of access roads into the littoral
forest for mineral explorations, for example, as in Petriky
between 1984 and 1992 (Ingram & Dawson 2006). This
is the case for the whole of Madagascar where deforestation is closely related to the expansion of roads and
accessibility (Chomitz et al. 2007). It has also been suggested that QMM’s (mining company) newly established
presence in the littoral forest since the 1990s has made
land tenure rights by local Anosy communities more uncertain and contested (Ingram & Dawson 2006), leaving
the littoral forest as open access for other groups to exploit. It is, however, beyond the scope of this study to
delve deeper in the different interpretations of deforestation and the local social institutions context needed to
protect scare resources. What we do know from a study
of socio-ecological systems in southern Madagascar is that
the institutional context and conditions of maintained
and well-defined property rights are important for maintaining forest cover (Elmqvist et al. 2007).
Ecosystem management
Southeast Madagascar is a rich biological and cultural
landscape containing many highly biodiverse forest types.
The littoral forest is among these and has exceptional
conservation and social value. If a long-term ecological
perspective is included in this picture, it becomes apparent that the littoral forest fragments are highly resilient
to climate change in spite of being naturally fragmented,
and that they have acted as refuges and centers of diversification over the past few millennia. These findings
have far-reaching implications and suggest that such ancient forest fragments may act as refuges for less-mobile
Conservation Letters 2 (2009) 163–170
c
Copyright and Photocopying: 2009
Wiley Periodicals, Inc.
Ecosystem management in Madagascar
M. Virah-Sawmy
organisms and as stepping-stones for others during predicted anthropogenic climatic change. It is therefore of
paramount importance to integrate the littoral forest and
other ancient fragments within a network of protected
areas.
This perspective provides evidence for a modified version of the biogeographic model suggested by Wilmé et al.
(2006) for microevolution in Madagascar, which predicts
several centers of endemism in the lowland and coastal
parts of the island. They hypothesized that river catchments with low-elevation sources were zones of isolation during climatic desiccation because organisms could
not find refuge along riverine corridors to forest refugia on higher elevation, leading to extensive speciation
of coastal taxa. By contrast, long-term ecological data
from the mid-Holocene indicate that the littoral forest in
the southeast still maintained biodiversity in small pocket
refuges during pronounced climatic desiccation (VirahSawmy et al. 2009, in press). Possibly, it is the isolation and reconnection processes to forest refuges caused
by differential sensitivity and resilience of plant assemblages to paleoclimatic variation that might play an important role in speciation. We suggest that additional siteselection criteria for conservation planning under global
climate change need to be considered such as interactions
between forest and freshwater systems that maintain soil
moisture during prolonged drought periods.
Mining and local demand for charcoal are threatening the survival of the littoral forest. Whilst the demands
for charcoal are being addressed regionally through exotic forest plantations, the impact of mining on biodiversity remains problematic. Increasingly, mining companies
such as Rio Tinto in Madagascar are involved in off-sites
biodiversity offsets to compensate for residual, unavoidable harm to biodiversity. While the benefits of biodiversity offsets are potentially large and will play an important role in responsible mining, several hurdles need to
be crossed to achieve them (Bishop et al. 2008). Questions
need to be answered on the ratio of off-site offset to onsite restoration that is acceptable in the case of converted
habitats that have high or even unique conservation and
social values. Further, should developers offset their indirect impacts (e.g., impacts arising from labor migration,
construction of roads)? Based on our analysis, we suggest
that if the appropriate conditions and measures are put
in place for restoration, for example, maintaining keystone species and ecological processes and the hydrological cycle, the littoral forest may once again expand and
regenerate after the degradation it suffered in the last few
decades, and it may retain its resilience to survive other
episodes of climatic instability.
Finally, there is great need for ecologists to understand the integrated nature of ecological, geomorpholog-
ical, climatical, and social systems to help the challenges
of responsible land-use with the maintenance of critical
biodiversity and ecosystem services. Cross-fertilization of
long-term with contemporary and inter-disciplinary perspectives has much to offer for conservation strategies in
times of global change. Importantly, the valuable lessons
learnt from this conservation hotspot have implications
for many other forests of the world’s that are coming under increasing pressure from the combined effects of climate change and land-use change.
Acknowledgment
Special thanks to J. Ebeling, L. Gillson, K. Willis, C. Bradshaw, A. Rodrigues, and two anonymous reviewers for
helpful comments on this manuscript. I would like to
thank the Oxford University Centre for the Environment, Wingate Foundation, Rufford Small Grant, Environment Change Institute, and Jesus College for financial
help.
References
Bishop, J., Kapila S., Hicks F., Mitchell P., Vorhies F. (2008)
Page 159 in Building biodiversity business. Shell International
Limited and the International Union for Conservation of
Nature, London and Gland.
Burney, D.A. (1987) Pre-settlement vegetation changes at
lake Tritrivakely, Madagascar. Palaeoecology of Africa and the
Surrounding Islands 18, 357–381.
Chomitz, K.M., Buys P., De Luca G., Thomas T.S.,
Wertz-Kanounnikoff S. (2007) At loggerheads?: Agricultural
expansion, poverty reduction, and environment in the tropical
forests. World Bank Policy Research Report, Washington,
D.C.
Consiglio, T., Schatz G.E., McPherson G., et al. (2006)
Deforestation and plant diversity of Madagascar’s littoral
forests. Conserv Biol 20, 1799–1803.
Cornet, A. (1974) Essai de cartographie bioclimatique à
Madagascar. ORSTOM, Paris, France.
de Gouvenain, R.C., Silander J.A. (2003) Littoral forest. Pages
103–111 in S.M. Goodman & J.P. Benstead, editors. The
natural history of Madagascar. University of Chicago Press,
Chicago, Illinois.
Dove, M.R. (2006) Indigenous people and environmental
politics. Annu Rev Anthropol 35, 191–208.
Du Puy, D.J., Moat J. (2003) Using geological substrate to
identify and map primary vegetation types in Madagascar
and the implications for planning biodiversity
conservation. Pages 51–74 in S.M. Goodman & J.P.
Benstead, editors. The natural history of Madagascar. The
University of Chicago Press, Chicago, Illinois.
Dufils, J.M. (2003) Remaining forest cover. Pages 88–96 in
S.M. Goodman & J.P. Benstead, editors. The natural history
c
Conservation Letters 2 (2009) 163–170 Copyright and Photocopying: 2009
Wiley Periodicals, Inc.
169
Ecosystem management in Madagascar
M. Virah-Sawmy
of Madagascar. The University of Chicago Press, Chicago,
Illinois.
Elmqvist, T., Pyykönen M., Tengö M., Rakotondrasoa F.,
Rabakonandrianina E., Radimilahy C. (2007) Patterns of
loss and regeneration of tropical dry forest in Madagascar:
the Social Institutional Context. PLoS ONE 5, 1–14.
Folke, C., Carpenter S., Walker B., et al. (2004) Regime shifts,
resilience, and biodiversity in ecosystem management.
Annu Rev Ecol Evol Syst 35, 557–581.
Ganzhorn, J.U., Lowry P.P., Schatz G.E., Sommer S. (2001)
The biodiversity of Madagascar: one of the world’s hottest
hotspots on its way out. Oryx 35, 346–348.
Goodman, S.M., Ramanamanjato J.-B. (2007) A perspective
on the paleo-ecology and biogeography of extreme
southeastern Madagascar, with special reference to
animals. Pages 25–48 in J.U. Ganzhorn, S.M. Goodman &
M. Vincelette, editors. Biodiversity, ecology and conservation of
littoral ecosystems in southeastern Madagascar,Tolagnaro (Fort
Dauphin). Smithsonian Institution, Washington, DC.
Green, G.M., Sussman R.W. (1990) Deforestation history of
the eastern rain forests of Madagascar from satellite images.
Science 248, 212–215.
Hannah, L., Dave R., Lowry P.P., et al. (2008) Climate change
adaptation for conservation in Madagascar. Biol Lett 4,
590–594.
Harper, G.J., Steininger M.K., Tucker C.J., Juhn D., Hawkins
F. (2008) Fifty years of deforestation and forest
fragmentation in Madagascar. Environ Conserv 34, 325–333.
Humbert, H., Cours Darne G. (1965) Notice de la Carte
Madagascar. Section Scientifique et Technique de L’Institut
Francais de Pondichéry, Pondichéry, India.
Ingram, J.C., Dawson T.P. (2006) Forest cover, condition, and
ecology in human-impacted forests, South-Eastern
Madagascar. Conserv Soc 4, 194–230.
Ingram, J.C., Whittaker R.J., Dawson T.P. (2005) Tree
structure and diversity in human-impacted littoral forests.
Environ Manage 35, 779–798.
Jarosz, L. (1993) Defining and explaining tropical
deforestation: shifting cultivation and population growth in
colonial Madagascar (1896–1940). Econ Geogr 69, 366–379.
Lowry, P.P.I., Schatz G.E., Phillipson P.B. (1997) The
classification of natural and anthropogenic vegetation in
Madagascar. Pages 93–123 in S.M. Goodman & B.D.
Patterson, editors. Natural change and human impact.
Smithsonian Institution Press, Washington, D.C.
170
MAP (2006) MADAGASCAR ACTION PLAN 2007–2012. A
bold and exciting plan for rapid development. In: Malagasy
Government, Republic of Madagascar.
Mittermeier, R.A., Gil P.R., Hoffman M., et al. (2005) Hotspots
revisited. University of Chicago Publishers, Chicago, Illinois.
Myers, N., Mittermeier R.A., Mittermeier C.G., da Fonseca
G.A.B., Kent J. (2000) Biodiversity hotspots for
conservation priorities. Nature 403, 853–858.
Noss, R.F. (2001) Beyond Kyoto: Forest management in a
time of rapid climate change. Conserv Biol 15, 578–590.
Pressey, R.L., Cabeza M., Watts M.E., Cowling R.M., Wilson
K.A. (2007) Conservation planning in a changing world.
Trends Ecol Evol 22, 583–592.
QIT Madagascar Minerals S.A. (2001) Pages 1–45 in Ilmenite
project: summary of social and environmental impact assessment.
QIT Madagascar Minerals, South Africa (QMM S.A.),
Montreal, Canada.
Vincelette, M., Théberge M., Randrihasipara L. (2007)
Evaluations of forest cover at regional and local levels in
the Tolagnaro Region since 1950. Pages 49–58 in J.U.
Ganzhorn, S.M. Goodman & M. Vincelette, editors.
Biodiversity, ecology and conservation of littoral ecosystems in
southeastern Madagascar,Tolagnaro (Fort Dauphin).
Smithsonian Institution, Washington, D.C.
Virah-Sawmy, M., Gillson L., Willis K.J. (in press) Does
heterogeneity enhance resilience to rapid climatic changes?
Sea-level rise, aridity and ecological dynamics in southeast
Madagascar. Ecology.
Virah-Sawmy, M., Willis K.J., Gillson L. (2009) Threshold
response of Madagascar’s littoral forest to sea-level rise.
Global Change Ecol Biogeogr 18, 98–110.
Warman, L., Moles A. (2009) Alternative stable states in
Australia’s wet tropics: a theoretical framework for the field
data and a field-case for the theory. Landscape Ecol 24, 1–13.
Willis, K.J., Birks H.J.B. (2006) What is natural? The need for
a long-term perspective in biodiversity conservation. Science
314, 1261–1265.
Willis, K.J., Gillson L., Virah-Sawmy M. (2008) Nature or
nurture: the ambiguity of C 4 grasslands in Madagascar. J
Biogeogr 35, 1741–1742.
Wilmé, L., Goodman S.M., Ganzhorn J.U. (2006)
Biogeographic evolution of Madagascar’s microendemic
biota. Science 312, 1063–1065.
Conservation Letters 2 (2009) 163–170
Editor: Dr. Ana Rodrigues
c
Copyright and Photocopying: 2009
Wiley Periodicals, Inc.