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Earth observation: overlooked potential to support species

reintroduction programmes
Terri P. Freemantle1, Tim Wacher2,3, John Newby3 and Nathalie Pettorelli1*
1
Institute of Zoology, Zoological Society of London, Regent’s Park, London NW1 4RY, U.K., 2Conservation Programmes (CP), Zoological Society of
London, Regent’s Park, London NW1 4RY, U.K. and 3Sahara Conservation Fund, Rue des Tigneuses 2, L’Isle, 1148, Switzerland

Abstract temporelle et spatiale de l’evaluation d’habitats. Cette


Habitat evaluation is considered an essential step for etude utilise des donnees d’observations terrestres libre-
assessing the potential for an area to support a viable ment accessibles pour evaluer si la Reserve de Faune
reintroduced population. Remote sensing techniques can d’Ouadi Rime-Ouadi Achim, au centre du Tchad, est
support such investigations, by greatly enhancing the propice  a la reintroduction de l’oryx algazelle. La reserve
temporal and spatial coverage of habitat assessments. This etait le dernier bastion de l’espece dans son aire de
study makes use of freely available earth observation data repartition historique avant son extinction  a l’etat sauvage
to evaluate the suitability of the Ouadi Rime–Ouadi Achim dans les annees 1980, ce qui donne un fondement a  cette
Game Reserve in central Chad for the reintroduction of the reintroduction. Les resultats montrent que, depuis les
Scimitar-horned Oryx. The reserve was the last stronghold annees 1980, il y a eu une forte augmentation des
of the species within its historical range, prior to extinction precipitations annuelles moyennes et des estimations de
in the wild in the 1980s, providing a basis for reintroduc- production primaire derivees des donnees satellites. La
tion. Results show that since the 1980s, there has been a reduction spatiale de la tendance moyenne de la produc-
steady increase in average annual precipitation and tion primaire montre que le nord de l’aire protegee
satellite-derived estimates of primary production. The presentait une tendance a  l’aridification entre 1982 et
spatial downscaling of the average trend in primary 2008 alors que le sud etait plut^  un reverdiss-
ot associe a
production showed that the north of the protected area ement intense. Par consequent, la zone de transition
exhibited a drying trend over the period 1982–2008, while subdesertique preferee des oryx est actuellement en voie de
the south was associated with intense greening. As a reduction. Si ceci est exact, cela signifie une possible
result, the subdesert transition zone preferred by oryx is reduction de l’habitat favorable aux oryx, ce qui pourrait
currently narrowing. If this is correct, this implies a avoir un effet negatif sur la reussite de l’etablissement
potential reduction of favourable habitat for the oryx, d’une population reintroduite autosuffisante.
which could have detrimental effects on the success of
establishing a self-sustaining reintroduced population.

Key words: earth observations, NDVI, oryx, remote sens- Introduction


ing, Sahara, translocations
Anthropogenic activities are having an increasingly neg-
ative impact on biodiversity at global, regional and local
Resume scales, and the resulting species loss is occurring at an
alarming rate (Pimm et al., 1995; Millennium Ecosystem
On estime qu’une evaluation de l’habitat est une etape
Assessment, 2005). In the face of habitat degradation,
essentielle pour estimer quel potentiel a une region de
increased resource exploitation, urbanization and devel-
supporter une population reintroduite viable. Les tech-
opment, alongside climate change and the associated
 distance peuvent aider lors de ces
niques de detection a
uncertainties involving global and local scale impacts, it is
investigations en augmentant fortement la couverture
globally recognized that we must preserve biodiversity to
secure our own well-being (Planet under Pressure, 2012).
*Correspondence: E-mail: Nathalie.Pettorelli@ioz.ac.uk

© 2013 Blackwell Publishing Ltd, Afr. J. Ecol. 1


2 Terri P. Freemantle et al.

One way to reduce biodiversity loss is to reintroduce species quantity and distribution) and its relationship to climate
where they have been extirpated. Reintroduction is defined as (Herrmann, Anyamba & Tucker, 2005) and species distri-
‘an attempt to establish a species in an area which was once part of bution and abundance (Pettorelli et al., 2011). When it
its historical range, but from which it has been extirpated or comes to support reintroduction efforts, approaches based on
become extinct’ (IUCN/SSC Re-Introduction Specialist Group, species distribution modelling need to be complemented with
1998). While reintroduction attempts are laudable and have information on long-term trends in ecosystem processes and
been numerous, their success rates have been generally low threats in potential reintroduction sites.
(Osborne & Seddon, 2012). For example, reviews of published This study proposes a way to overcome the problem of
literature on reintroductions identify success rates as low as conducting habitat assessments over large spatial extents
11% (Beck et al., 1994) and 26% (Fischer & Lindenmayer, and high temporal resolutions, aiming to promote a more
2000), although self-assessment by reintroduction practi- integrative use of EO products and to develop a low-cost
tioners can result in higher scores of success (Soorae, 2010). remote sensing-based framework for habitat assessments, in
The factors that impede the success of reintroductions have compliance with the IUCN Reintroduction Specialist Group
not been documented widely, but there is evidence that the Guidelines. It uses the case study of the Scimitar-horned
quality and quantity of suitable habitat is a primary driver of Oryx (Oryx dammah) and its extirpation from the Ouadi
success (Wolf et al., 1996; Armstrong et al., 2002; Cook, Rime–Ouadi Achim Game Reserve (OROAGR). This oryx
Morgan & Marshall, 2010; Osborne & Seddon, 2012). species is currently designated as Extinct in the Wild by the
Surprisingly, a large proportion of the reintroduction IUCN (2011). Fortunately, as a result of large-scale captive-
literature fails to document research into habitat quality breeding programmes, a large ex situ population now exists
and functioning prior to release of the population (but see dispersed widely across Europe, the Middle East and U.S.A.
Cook, Morgan & Marshall, 2010). Moreover, while the IUCN/ (Gilbert & Woodfine, 2008; JN & TW, pers. comm.). From this
SSC Re-Introduction Specialist Group (1998)guidelines state global population, it is possible to reintroduce the scimitar-
that habitat assessments prior to release are necessary, they horned oryx back to its native habitats, providing political
offer little advice as to how this should be achieved. As a result, and social support exist, and habitat quality and ecological
there is no standardized framework for habitat quality functioning are shown to be sufficient for successful
assessment. One can expect that if implemented effectively, reintroduction. Based on the latest distribution models for
such a framework could positively impact the success rate of this species, the OROAGR has been proposed as a potential
reintroductions. reintroduction site (Wacher et al., 2011a,b). Here, we
Traditional assessments of ecological properties have often address habitat quality and ecological function at OROAGR
been carried out using time-consuming and labour-intensive by using 27 years of satellite imagery. More precisely, we
field surveys, which have the disadvantage of potential assessed long-term trends in primary production and
inconsistence over space and time (Pfeifer et al., 2012). climatic conditions in this protected area, as well as
Satellite-based information, however, offers a way to assess potential spatial variation in anthropogenic threats. To
habitat type and condition over large areas. This information clarify the links between primary production, climatic
can be used in species distribution modelling exercises variation and anthropogenic pressure in the area and
(Lehmann, Overton & Austin, 2002; Scott, 2002), maxi- inform discussions on how future global environmental
mum entropy modelling (Phillips, Anderson & Schapire, change might impact the suitability of the OROAGR for
2006) and ecological-niche factor analysis (Hirzel et al., oryx, we investigated two primary hypotheses: H1) Precip-
2002) to identify areas best matching the niche require- itation is a main driver of long-term trends in primary
ments of the species to be reintroduced. These approaches, production in the area. H2) At a local scale, high anthro-
however, rely on the use of temporally static data and do not pogenic activity negatively impacts primary production.
make use of the full potential of earth observations to inform
reintroduction efforts. In recent years, access to spatial data
Material and method
has significantly increased, as has the spatial, temporal and
spectral resolution of satellite sensors (Pfeifer et al., 2012).
The Ouadi Rime–Ouadi Achim Game Reserve (OROAGR)
Earth observation (EO) data have been used in numerous
ecological studies of habitat functioning, providing, for The OROAGR is located in central Chad at 15.77˚N
example, information on vegetation dynamics (phenology, 19.00˚E (Fig. 1). The designated reserve area covers

© 2013 Blackwell Publishing Ltd, Afr. J. Ecol.


Earth observations and species reintroductions 3

Fig 1 Location of the Ouadi Rime–Ouadi


Achim Game Reserve (OROAGR) and
landcover class within the OROAGR
derived from GLOBCOVER (2010). Climate
averages for the OROAGR over the period
1950–2000 are derived from Worldclim

77,950 km² and local altitude ranges between 190 and addax (Addax nasomaculatus; CR), the endangered African
461 m (Keith & Plowes, 1997; Hartley et al., 2007). The hunting dog (Lycaon pictus) and other vulnerable species,
environment is typical of that associated with the Sahelian such as cheetah (Acinonyx jubatus). There is, however, no
sub-Saharan zone and is characterized by high average recent evidence for the presence of these species in the
temperatures and moderate/low annual rainfall (Fig. 1). area.
Three major habitat types can be distinguished from south The decline of scimitar-horned oryx in the OROAGR has
to north: 1) Sahelian wooded grasslands, 2) subdesert been documented in detail (Newby, 1980; East, 1990;
grassland and 3) desert, with subdesert grassland covering Wilson & Reeder, 1993). Because of the historical abun-
approximately two-thirds of the protected area (Hartley dance of scimitar-horned oryx, insufficient protection was
et al., 2007) (Fig. 1). Topographically speaking, the assigned to this species. This left it vulnerable to poaching
reserve maintains a relatively featureless low terrain. (Bassett, 1975; Newby, 1980) and to exclusion from good
However, the area that borders in the east is a series of quality pasture by the presence of nomadic herdsmen and
massifs reaching approximately 1500 m, from which their herds (Bassett, 1975). The oryx were traditionally
arises a network of east-to-west flowing wadis. These hunted for their meat and hides. Civil war and the
significantly enhance the biological diversity of the reserve introduction of automated weapons and motorized vehi-
as a result of their associated temporary pools, floodplains cles increased the hunting pressure on the species,
and inundation zones, providing a source of freshwater. resulting in a devastating decline in the OROAGR popu-
The term ‘Sahel’ in Arabic literally translates as ‘shore’ lation (Newby, 1980). At a larger scale, overhunting and
and refers to the ‘sparsely vegetated fringe of sand seas’ habitat loss, including competition with domestic livestock,
(Newby, 1980): as well as being one of Africa’s largest have been reported as reasons for the extinction of many
protected areas, the OROAGR is also one of the only wild oryx populations (Mallon & Kingswood, 2001;
designated protected areas that falls in the transition zone Beudels et al., 2006; Morrow, 2013).
between true desert (Sahara) and Sahel.
The OROAGR was gazetted in 1969 and was, until the
Earth observation data
early 1980s, known as a stronghold for the scimitar-
horned oryx. The protected area is of significant ecological The Normalized Difference Vegetation Index (NDVI) was
importance for a number of factors, primarily the presence used to monitor spatial and temporal trends in primary
of IUCN classified fauna, such as the critically endangered production (Pettorelli et al., 2005). The NDVI is amongst
(CR) dama gazelle (Gazella dama) and the largest known the most intensely studied and commonly used vegetation
population of vulnerable dorcas gazelle (Gazella dorcas) indices (Pettorelli et al., 2011). It is derived from the red:
(IUCN, 2011). The area was also historically home to the near-infrared reflectance ratio (NDVI = (NIR-RED)/

© 2013 Blackwell Publishing Ltd, Afr. J. Ecol.


4 Terri P. Freemantle et al.

(NIR + RED), where NIR and RED are the amounts of 2002; Leroux et al., 2010) was used to assess the human
near-infrared and red light, respectively, reflected by the impact in the study area. The Human Footprint Index
vegetation and captured by the sensor of the satellite (HFI) data were acquired from the Socio-Economic Data
(Jensen, 2007). NDVI data varies from 1 to +1: NDVI and Applications Centre (http://sedac.ciesin.org/wildare-
values below 0 are associated with an absence of as/); its native spatial resolution is 1 km. The map
photosynthetically active vegetation. NDVI data were identifies, and rates on a scale of 1–100, anthropogenic
obtained from the Global Inventory Modelling and Map- impacts on the environment. A rating of 1 indicates
ping Studies (GIMMS; Tucker et al., 2005) Advanced Very minimal impact from humans and 100 high impacts. HFI
High Resolution Radiometer. The GIMMS product pro- is a unit-less index.
vided NDVI data at 8-km resolution for the period 1981–
2008 (http://www.glcf.umd.edu/data/gimms/), giving a
Analysis
total time series of 27 years. The native temporal fre-
quency of the GIMMS product is 15 days. In sparsely The protected area boundary of the OROAGR was obtained
vegetated areas such as the one considered here, the from the World Database of Protected Areas (downloaded
relationship between NDVI and primary production can be in February 2009). Because conditions outside a given
biased and the use of soil-adjusted vegetation indices such protected area can sometimes significantly differ from
as SAVI has been previously recommended (Huete, 1988). conditions inside that area (Hartley et al., 2007), a 30-km
However, SAVI requires local calibration and such infor- buffer was created for the OROAGR, and relevant envi-
mation was not available for our area. Moreover, there is ronmental data were extracted for the buffer as well as for
currently no processed time series available for SAVI (or the protected area (Pettorelli et al., 2012).
any other soil-adjusted index) for the period considered. To correct for environmental noise (smoothing) in the
Because we focused our analyses on the search for extracted NDVI data (Pettorelli et al., 2005), rapid
significant changes in NDVI over time, our conclusions decreases (or ‘drops’) in NDVI values (of 0.25 or more
are not expected to be significantly affected by the use of from one composite to the next) that were immediately
NDVI over SAVI. followed by a return to the original values or higher were
Precipitation data were acquired from WorldClim identified for each pixel (Garonna et al., 2009). We
(wordclim.org) and the Global Precipitation Climatology attributed these changes to environmental noise, because
Project (GPCP; Herrmann, Anyamba & Tucker, 2005; factors such as water, clouds and shadow give rise to
Gruber & Levizzani, 2008) v2.2. WorldClim provided highly reduced NDVI values (Pettorelli et al., 2005). Once
historical climate averages (Hijmans et al., 2005). The we had identified all contaminated values, they were
GPCP data provided monthly estimated values of precip- replaced by the average of the previous and following
itation at the global scale and at a 2.5 degree global grid values, so as to ‘smooth’ the annual NDVI curve for that
resolution; these data spanned from 1979 to present day. pixel. If two consecutive ‘drops’ values were present, the
Landcover raster data sets were acquired from the most average of the closest higher NDVI values was calculated;
recent release of GLOBCOVER (2010). The GLOBCOVER all pixels with 3 consecutive NDVI values <0 were removed
project was initiated in 2004 by the European Space (Garonna et al., 2009).
Agency to document and produce a land cover map using Four NDVI-based parameters were then calculated, all
300-m data primarily obtained from the MERIS sensor on representing vital indicators of ecosystem functioning
board ENVISAT. The newly released data were obtained (Alcaraz-Segura et al., 2009; Pettorelli et al., 2012):
for 2009–2010 (http://ionia1.esrin.esa.int/). The landcov- annual NDVI maximum (MAX), annual NDVI minimum
er composites have been composed using data from the (MIN), integrated-annual NDVI (I-NDVI) and annual
MERIS sensor and corrected for cloud cover, atmospheric relative range [RREL = (MAX MIN)/MEAN, where MEAN
effects, geo-localization and remapping. The classification is the annual mean NDVI]. These four parameters were
used is compatible with the United Nation Land Cover estimated at the protected area and at the GIMMS NDVI
Classification system. pixel scale (i.e. the four parameters were determined for
The human footprint, which is a composite of transport, each of the 1304 pixels within the protected area). The
night-time lights, urban areas, land cover and population I-NDVI corresponds to the sum of positive NDVI values
density layers, normalized by biome (Sanderson et al., over a set period in time, in this instance calculated as the

© 2013 Blackwell Publishing Ltd, Afr. J. Ecol.


Earth observations and species reintroductions 5

sum of all NDVI values over a year. This alongside Table 1 Modelling spatial correlation in I-NDVI at the NDVI pixel
maximum value of NDVI has been shown to be a good scale (n = 1304 NDVI pixels located in the Ouadi Rime–Ouadi
measure of overall production and biomass (Reed et al., Achim Game Reserve). Four model structures (Null, Gaussian,
Exponential and Spherical) were considered to determine the form
1994). Minimum NDVI was then used alongside the other
of the spatial autocorrelation in I-NDVI. Based on both Akaike
parameters to determine intra-annual relative range,
Information Criterion (AIC; Akaike, 1974) and Bayesian Infor-
which provides information about the level of annual mation Criterion (BIC; Schwarz, 1978), the data were found to
seasonality in greenness dynamics (i.e. the lower the RREL, best fit an exponential distribution
the lesser seasonal the environment is).
Model structure AIC BIC
All statistical analyses were performed in the statistical
package R (www.r-project.org). To assess the existence or 1: Null 8200.70 8231.72
nonexistence of significant temporal trends in these four 2: Gaussian 35,906.09 35,942.28
3: Exponential 6908.91 6945.10
parameters over the period 1982–2008 at both scales,
4: Spherical 6925.10 6961.29
Mann–Kendall trend tests were used (Pettorelli et al.,
2012). For some pixels, the level of contamination was
high and smoothing was not possible. For these pixels, it
was therefore not possible to directly assess the existence or 8.39, while being estimated at 9.15 in a 30-km buffer,
nonexistence of significant temporal trends in these four indicating low levels of human activity in the area.
parameters over the period 1982–2008. To tackle this Dominant land cover classes and percentage area cover
issue, correlations were spatially interpolated to estimate within the protected area were calculated as bare areas
the likely significance of any temporal trend in any of the (77.79%), closed to open grassland (21.95%), sparse
four parameters considered for highly contaminated pixels; vegetation (0.15%), mosaic forest-shrublands/grass
this was done using inverse distance weighting (Shepard, (0.09%) and mosaic vegetation/croplands (0.004%). Sim-
1968). ilarly for the 30-km buffer zone, bare areas (65.9%), closed
To explore the relationship between the long-term to open grassland (33.01%), sparse vegetation (0.21%),
average I-NDVI, human impact and climate in the mosaic forest-shrublands/grass (0.53%), rain fed croplands
OROAGR, an 8 9 8 km reference grid was overlaid on (0.15%), mosaic vegetation/croplands (0.07%), and
top of the HFI and GPCP precipitation data sets. Values mosaic cropland/vegetation (0.02%).
within each reference grid cell were extracted and aver- Mann–Kendall tests revealed that there was a positive,
aged to give a corresponding value for each pixel at the yet nonsignificant, long-term trend in I-NDVI for the
same resolution as the GIMMS NDVI data. Observations protected area (τ = 0.248, n = 27 years, P = 0.07). Our
closer together will tend to be more similar to those further analyses also revealed a significant increase in annual
apart (Cressie, 1991), leading to spatial autocorrelation. MAX and RREL in the reserve (MAX: τ = 0.419,
To assess the effects of human impact and average n = 27 years, P = 0.002; RREL: τ = 0.436, n = 27 years,
precipitation on I-NDVI while correcting for this source P = 0.001). However, a significant decrease in MIN NDVI
of bias, Laird–Ware linear mixed models were used (Laird was observed in the protected area (τ = 0.282,
& Ware, 1982). These allow the direct incorporation of the n = 27 years, P = 0.04). Similar results were obtained
spatial autocorrelation structure in the residuals for the when considering NDVI data from the buffer zone (see also
considered models. The spatial autocorrelation structure Figs 2 and 3). The long-term trends in the four NDVI-
was assessed using semi-variogram: four possible struc- based parameters considered were concomitant with the
tures (Null, Gaussian, Exponential and Spherical) were observation of a significant increase in annual average
compared using the Akaike Information Criterion (AIC; precipitation over the period 1979–2011 (τ = 0.274,
Akaike, 1974; Table 1). n = 32 years, P = 0.03; Fig. 2). As expected (H1), annual
variation in I-NDVI in the OROAGR were significantly
correlated to changes in annual average precipitation in
Results
the reserve (slope = 0.036  0.0072, n = 27, R² = 0.47,
The long-term average annual precipitation for OROAGR P < 0.001).
is 77 mm with an average annual temperature of 29˚C Over the period 1982–2008, only 17.7% of the OROAGR
(Fig. 1). Average human footprint in the reserve reached exhibited a significant increase in I-NDVI, while 28.4% of the

© 2013 Blackwell Publishing Ltd, Afr. J. Ecol.


6 Terri P. Freemantle et al.

Fig 2 Relationship between the I-NDVI


and average precipitation for the Ouadi
Rime–Ouadi Achim Game Reserve (ORO-
AGR). Trends in I-NDVI are provided (a)
for the OROAGR and (b) for the pixels
located in the 30-km buffer around the
OROAGR

OROAGR was associated with significant decreases in this 2005; Herrmann, Anyamba & Tucker, 2005), both of
parameter (Table 2). Similar results were reported for all four which documented a steady increase in precipitation since
parameters (Table 2, Fig. 3). Based on (H2), we expected severe droughts occurred in the region in the 1970s. These
spatial variation in anthropogenic activity to negatively increases in precipitation have been interpreted as being
impact I-NDVI in the reserve: we could not report any indicative of a ‘postdrought recovery’ caused by precipita-
significant influence of HFI on the long-term I-NDVI average tion in the region returning to normal conditions at the
for the OROAGR (Table 3). At such scale (i.e. at the OROAGR end of the drought period. Other causes, however, can be
NDVI GIMMS pixel scale), only changes in precipitation considered, such as feedback from climate change which is
regimes were reported to significantly influence long-term predicted to increase precipitation in the west of Africa
averages in I-NDVI values (slope = 65.78  3.93, (Boko et al., 2007).
n = 1304 years, P < 0.001; Fig. 4). An important result from our study is the increasing
trend in net primary production over the past 27 years at
the scale of the reserve. However, what are the factors
Discussion
driving these reported patterns in primary production
Habitat assessments are an invaluable tool when dynamics for this area? There are several possibilities:
considering management strategies for species reintro- Herrmann, Anyamba & Tucker (2005) propose, while
ductions. Remote sensing can provide essential informa- rainfall is often considered the dominant cause for
tion on long-term changes in climatic conditions and increased greenness, there is evidence of another possible
primary production, which can help assess how suitable cause, namely the existence of a human-induced land
an area might be for reintroduced species. Our results cover change and anthropogenic activities. Nemani et al.
show that (1) since the 1980s, there has been a steady (2003) also discuss the possibility that climate-driven
increase in precipitation and satellite-derived estimates of global increases in primary production between 1982 and
primary production; (2) precipitation regime is the main 1999 could be the result of increased carbon dioxide
driver of vegetation dynamics in this area; (3) anthro- improving plants photosynthetic functioning, a direct
pogenic impact in the region, as derived from EO data, is response to climate change. However, when testing the
low; (4) the long-term trends in vegetation dynamics theories of Herrmann, Anyamba & Tucker (2005) regard-
vary spatially, with the north of the protected area ing human impact on net primary production, we found
exhibiting a drying trend over the period 1982–2008, no evidence to support this hypothesis (H2). This can be
and the south an intense greening. explained by the low incidence of average human impact
Our report of increasing precipitation in the OROAGR in the region (here indexed by land cover and human
over the last three decades is supported by previous studies footprint). Furthermore, investigation into the temporal
undertaken in the Sahel region (Anyamba & Tucker, trends in I-NDVI values in the OROAGR identifies a

© 2013 Blackwell Publishing Ltd, Afr. J. Ecol.


Earth observations and species reintroductions 7

Fig 3 Spatial variation in vegetation


dynamics parameters within the protected
area. spatial patterns in integrated-annual
NDVI (I-NDVI), intra-annual relative
range (seasonality – RREL), maximum
and minimum NDVI (MAX and MIN) are
provided (a) for the OROAGR and (b) for
the pixels located in the 30-km buffer
around the OROAGR

response to El Nino/Southern Oscillation (ENSO) events. A OROAGR? Higher I-NDVI values have been previously
significant drop in I-NDVI was observed during 1993– linked to change in herbivore abundance in the Kalahari,
1994, corresponding to a known El Nino year, while peaks Botswana (Verlinden & Masogo, 1997). Pettorelli et al.
were observed in 1988–1989, and 1999–2000 (Fig. 2) (2009) also found that, generally, NDVI estimates of
corresponding to known La Nina years (based on the primary production were positively correlated to species
Multivariate ENSO Index; NOAA, 2012). These observed abundance (based on 13 species of African ungulate).
patterns linking I-NDVI with ENSO events reiterate the Additionally, when considering the effect of NDVI on
notion that climate is the main driver of vegetation grassland ungulates, Mueller et al. (2008) found that
dynamics in the region. NDVI was a good predictor of gazelle presence in the arid
How can the reported trends be expected to influence the steppes of Mongolia. Increased I-NDVI values in the
success of a potential reintroduction of oryx in the OROAGR can thus be expected to correlate with increased

© 2013 Blackwell Publishing Ltd, Afr. J. Ecol.


8 Terri P. Freemantle et al.

Table 2 Percentage of pixels showing change in vegetation higher competition between domestic and wild species for
dynamics within the protected area for each NDVI-based resources. Moreover, unless the protection of the OROAGR
parameter is strictly enforced, increased concentrations of people in
Change I-NDVI RREL MAX MIN the reserve could lead to increased pressure (e.g. poaching)
on existing and reintroduced wildlife.
Significant decrease 28.37 2.33 30.87 23.42
Nonsignificant decrease 24.18 23 8.16 64.98 Spatial variation in greenness levels could then restrict
Nonsignificant increase 31.74 27.90 23.61 11.35 the success rate of a potential reintroduction, despite this
Significant increase 15.71 46.77 37.36 0.25 overall increase in food abundance. Our observations indeed
identify a trend whereby the north–south gradient of the
where MAX – annual NDVI maximum; MIN – annual NDVI region’s ecosystem is becoming more pronounced, and the
minimum; I-NDVI – integrated-annual NDVI; RREL – annual
transition zone (subdesert Steppe) between true Sahara and
relative range (RREL=[MAX-MIN]/MEAN, where MEAN is the
Sahelian Grassland is disappearing, or being ‘squeezed’ by
annual mean NDVI for the OROAGR).
the two subecosystems on either side. This ‘ecosystem
squeezing’ is evident in the increased seasonality (as indexed
Table 3 Satellite-based estimates of annual primary production in by the intra-annual relative range; RREL) in the region,
the Ouadi Rime–Ouadi Achim Game Reserve (as indexed using
indicative of larger variations between the MIN and MAX
I-NDVI), modelled as a function of precipitation and human
NDVI, and through the north’s substantial decrease in
footprint index (HFI)
I-NDVI compared with the south’s increase. Yet, as docu-
Parameter Slope Standard Error t value P value mented by Newby (1978, 1988), the oryx are primarily a
Intercept 46.38 2.58 17.96 <0.001 grassland and subdesert species, whose distribution is
HFI 0.21 0.16 1.28 0.20 dictated by available habitat and pasture. Oryx dietary
Precipitation 63.44 4.53 13.99 <0.001 habits are varied, though have been documented to favour
HFI 9 Precipitation 0.31 0.25 1.22 0.22 vegetation which thrives in the previously broad subdesert
transition zone, characterized by annual rainfall between
75 and 400 mm (Gilbert, 2004). As a result, species
distribution has been largely ‘concentrated to the northern
and southern subdesert fringes of the Sahara (Wakefield,
Engel & Gilbert, 2004). Our results suggest this subdesert
transition zone is diminishing in size. If this is correct, this
could ultimately result in a potential reduction of favourable
habitat for the oryx, which could have detrimental effects on
the success of establishing a self-sustaining reintroduced
population. More research is thus required to explore (a)
how the reported spatiotemporal patterns in vegetation
dynamics correlate with changes in land cover and land use;
(b) how these changes are likely to impact the distribution of
favourable habitats for oryx; (c) to which extent oryx will be
Fig 4 Relationship between vegetation dynamics and precipita-
able to adapt to the new spatial distribution of environmen-
tion in the Ouadi Rime–Ouadi Achim Game Reserve (OROAGR), at
the NDVI pixel scale (n = 1304 pixels). The I-NDVI is the sum of tal conditions occurring in the OROAGR.
all NDVI values over a set period of time; this correlation shows The availability of satellite data has enabled us to gain a
the I-NDVI and average precipitation calculated on a pixel-by- better understanding of the changes happening in the
pixel basis for the OROAGR world around us on a much larger scale than has ever
been possible before. With increased availability of higher
food availability for herbivorous ungulates in the area, resolution data, already known and established methods in
potentially positively impacting distribution and abun- remote sensing and spatial analysis can only be further
dance. But this greening trend may also lead to increased improved, with higher temporal, spatial and spectral
concentrations of livestock and people in the area, which frequency data. In the case we considered here, the use
could mean increased grazing pressure by livestock and of earth observation products has provided information on

© 2013 Blackwell Publishing Ltd, Afr. J. Ecol.


Earth observations and species reintroductions 9

ecosystem functioning with high spatial and temporal Alcaraz-Segura, D., Cabello, J., Paruelo, J.M. & Delibes, M. (2009)
frequency, something which could not have been achieved Use of descriptors of ecosystem functioning for monitoring a
by traditional ground surveying. These results have national park network: a remote sensing approach. Environ.
Manage. 43, 38–48.
provided a preliminary insight into the nature of ecosystem
Anyamba, A. & Tucker, C. (2005) Analysis of Sahelian vegetation
functioning within the reserve and have provided a dynamics using NOAA-AVHRR NDVI data from 1981–2003. J.
starting point to discuss future implications for the Arid Environ. 63, 596–614.
reintroduction of the scimitar-horned oryx. They have Armstrong, D.P., Davidson, R.S., Dimond, W.J., Perrott, J.K.,
also highlighted the need for further research into how Castro, I., Ewen, J.G., Griffiths, R. & Taylor, J. (2002)
spatial variability in net primary production could impact Population dynamics of reintroduced forest birds on New
a reintroduced population of oryx. Overall, our study thus Zealand islands. J. Biogeogr. 29, 609–621.
Bassett, T. (1975) Oryx and addax in Chad. Oryx 13, 50–51.
supports the notion that the use of earth observation
Beck, B.B., Papaport, L.G., Stanley Price, M.R. & Wilson, A.C.
products can greatly enhance the practise and manage-
(1994) Reintroduction of captive born animals.In: Creative
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icantly to their success. Admittedly, such a contribution animals(Eds P. J. S. Olney, G. M. Mace and A. T. C. Feistner).
might not be possible in all situations: NDVI is a relatively Chapman & Hall, London, U.K.
crude means of monitoring changes in resource availabil- Beudels, R.C., Devillers, P., Lafontaine, R-M., Devillers-
ity for herbivores, and information about changes in NDVI Terschuren, J. & Buedels, M-O.(2006) Sahelo-Saharan
Antelopes. Status and Perspectives. Report on the conservation
dynamics over time might be more useful for some species
status of the six Sahelo-Saharan Antelopes.In: CMS SSA
and some ecosystems than others (Pettorelli et al., 2011).
Concerted Action. 2nd edn. CMS Technical Series Publication N
Similarly, changes in specific seasons might be more °11, 2005. UNEP/CMS Secretariat, Bonn, Germany.
important than others: in 2000, Illius and O’Connor, for Boko, M., Niang, I., Nyong, A., Vogel, C., Githeko, A., Medany, M.,
example, demonstrated how ungulate population dynam- Osman-Elasha, B., Tabo, R. & Yanda, P. (2007) Contribution of
ics in arid and semi-arid grazing systems were particularly Working Group II to the Fourth Assessment Report of the
sensitive to the abundance of resources during the dry Intergovernmental Panel on Climate Change. In: Climate
season, as compared with that of the wet season (Illius & Change: Impacts, Adaptation and Vulnerability. Cambridge
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O’Connor, 2000). Tailoring the type of information that
Cook, C.N., Morgan, D.G. & Marshall, D.J. (2010) Reevaluating
can be extracted from satellite-based data to the ecosystem suitable habitat for reintroductions: lessons learnt from the
and biology of the species considered will thus be an eastern barred bandicoot recovery program. Anim. Conserv. 13,
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remote sensing data to reintroduction success. Cressie, N. (1991) Statistics for Spatial Data. John Wiley and Sons,
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Thanks are extended to the Sahara Conservation Fund
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(SCF), His Highness Sheikh Mohammed bin Zayed Al 1–11.
Nahyan, Crown Prince of Abu Dhabi, Emirates Center for Garonna, I., Fazey, I., Brown, M.E. & Pettorelli, N. (2009) Rapid
Wildlife Propagation (ECWP), International Fund for primary productivity changes in one of the last coastal
Houbara Conservation (IFHC), Conservation Programmes, rainforests: the case of Kahua, Solomon Islands. Environ.
Zoological Society of London, and finally the Institute of Conserv. 36, 253–260.
Zoology, for funding this research. Personal thanks are Gilbert, T. (2004) European studbook for scimitar-horned oryx
(Oryx dammah). Winchester, U.K.
also extended to Alienor Chauvenet for her technical
Gilbert, T. & Woodfine, T. (2008) The reintroduction of scimitar-
assistance. Additionally, thanks to James Duffy and Wil-
horned oryx Oryx dammah to Dghoumes National Park. Tunisia
liam Cornforth for their technical support and comments. Report to members of the European Endangered Species
Programme for scimitar-horned oryx.
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