Impacts of climatic changes on small mammal

communities in the Sahel (West Africa) as

evidenced by owl pellet analysis
Massamba Thiam1,2, Khalilou Bâ1 & Jean-Marc Duplantier3*
1
IRD, CBGP (UMR 022: IRD/INRA/CIRAD/SupAgroMontpellier), Centre ISRA/IRD de Dakar-Bel-Air,
BP 1386, Dakar CP 18524, Sénégal
2
Département de Biologie Animale, Université Cheick Anta Diop, BP 5005, Dakar, Sénégal
3
IRD, CBGP( UMR 022: IRD/INRA/CIRAD/SupAgroMontpellier), Campus International de Baillarguet,
CS 30016, 34 988 Montferrier sur Lez cedex, France
Received 16 April 2007. Accepted 4 March 2008

To evaluate the impact of climatic change on rodent sahelian communities, we analysed the
contents of over 2500 barn owl (Tyto alba) pellets collected along the Senegal river between
1989 and 2003, and from the Ferlo sahelian area in 2003. These results are compared with data
from the 1970s and 1980s in the same zones. Rodents were the most common prey (over 90%).
Gerbillinae were most common in dry areas (84 to 96%) whereas in wetlands and rice fields
murines were most common (77 to 88%). Nowadays, the genus Gerbillus constitutes the main
prey in dry areas (77% to 88%). The genus Taterillus, which was the most abundant rodent in
the Ferlo in the 1970s, now represents only 7% of rodents. Gerbils were not present in Senegal
before the 1980s: G. tarabuli and G. henleyi were trapped for the first time in 1989 at the north-
ern border of Senegal, and G. nigeriae 10 years later at the same place. The latter is now pres-
ent a hundred kilometres southwards and as abundant in owl pellets as the two other gerbils.
Key words: climatic change, Sahel, rodents, owl pellets, Gerbillus.

INTRODUCTION xerophylic elements as a consequence of the

The Sahel is a vast transitional zone between
the Sahara desert to the north and the arboreal
savannas and open forests of Sudan to the south.
It is thus a contact zone between two different
environments. Popov (1996) indicated that these
zones are characterized mainly by their ecological
instability and their tendency to undergo rapid
changes, particularly linked to meteorological
fluctuations. The strong climatic variations that
have occurred in the Sahel over the past decades
has been the focus of much research on this region.
Over the past centuries (Nicholson 1978) this zone
experienced considerable climatic changes, and
numerous authors have agreed that an obvious
desertification process has been under way since
the end of the 1960s over the entire Sahelian and
Sudanian zones in West Africa (see reviews by
Hulme 1992; Hulme et al. 1999, 2001; and Moron
1994). This desertification (Popov 1996) is most
striking in the arid part of the Sahel, where the
Sahara desert has progressed southwards. The
consequences of this can be clearly seen even in
the savanna, where certain plant species have
disappeared or have been replaced by more
*Author for correspondence. E-mail: duplanti@mpl.ird.fr
African Zoology 43(2): 135–143 (October 2008)
persisting lack of rain, leading to profound
changes in plant communities. In northern
Senegal, Handschumacher et al. (1992) indicated
that rainfall has been decreasing since the 1930s,
and that a marked drought began in the early
1970s. A comparison of the isohyets before and
after 1970 (Fig. 1), clearly shows that rainfall de-
creased by about 200 mm over the entire country
between 1947–1969 and 1970–1992 (Trape et al.
1996).
While there are numerous studies regarding
invertebrates (e.g. see Maleque et al. (2006),
Rombke et al. (2005) and Hodkinson (2005) for
recent references) and plants (see Goetze et al.
(2006) for recent fluctuations in the forest–savanna
mosaic of West Africa) as indicators for recent
global changes, only few studies have been
devoted to vertebrates (yet see Drapeau et al.
(2003) for birds and Whitfield & Elliot (2002) for
fishes). Although certain plants and invertebrates
react almost immediately to such modifications,
and may thus be used as indicators of temporary
variations, small vertebrates, and rodents in
particular, can be good indicators of more perma-
nent changes, since their capacity for dispersal is
136 African Zoology Vol. 43, No. 2, October 2008
Fig. 1. Location of study sites and comparison of isohyets in Senegal between 1947 and 1969, and 1970 and 1992,
(from Trape et al. 1996).
limited and they have high reproductive and pop-
ulation turnover rates. We thus hypothesized that
the climatic changes that have been proceeding
over the last decades have modified resident
rodent communities in the Sahel zone, based on a
longitudinal analysis of the prey found in regurgi-
tation pellets from barn owls in northern Senegal.
This method was chosen because owl pellets
provide easy access to information about the
species composition of rodent communities in a
given environment (Taylor 1994; Yom-Tov & Wool
1997). The most useful data can be obtained from
barn owl, Tyto alba, pellets (Saint-Girons & Spitz
1966). The prey remains in barn owl pellets provide
important and sometimes surprising information
(for species that are not easily trapped) regarding
the systematic composition and distribution of
particularly the small mammal and bird faunas
(Heim de Balsac 1965).
The objective of our study was to measure the
impact of the desertification in the Sahel on the
composition and status of small mammal (particu-
larly rodent) communities. To do this, we com-
pared the results from our recent collections (1990
to now), carried out in the Ferlo (Sahelian area in
Senegal), with data collected by Poulet (1972a,b,
1974, 1982, 1984) in the same biogeographical zone
during the 1970s. We examined in detail the genus
Gerbillus, which appeared in Senegal at the end of
the 1980s (Duplantier et al. 1991) and has been
expanding since (Bâ 2002).
MATERIALS & METHODS
Collecting localities
Between 1989 and 2003, during various expedi-
tions in northern Senegal, we collected regurgi-
tated pellets from barn owls in the district of
Richard-Toll, which lies 105 km from the town of
Saint-Louis on the left bank of the Senegal River
(16°27’N, 15°42’W). It is part of the Sahel region,
which is the hottest and most arid part of the
country, characterized by a long dry season lasting
8–9 months (November to June or July) and a
short rainy season lasting 3–4 months (Leroux
1983). Between 1951 and 1969, average annual
rainfall at the nearest official meteorological sta-
tion (Dagana, 20 km east) was 328 mm (S.D. = 98;
min/max = 152/630) , while between 1970 and 2000
it was only 197 mm (S.D. = 80; min/max = 58/328).
Maximum rainfall between 1970 and 2000 approxi-
mated only the average annual rainfall for the
period 1951–1969. Our samples were taken from
the rice fields at Colonat-Richard-Toll, and from an
airfield located east of Richard-Toll. The rice fields
lie in the humid Walo floodplain of the Senegal
 Thiam et al.: Impacts of climate changes on Sahel small mammal communities
River, which is temporarily flooded during the
rainy season; the airfield lies in the dry Diéry zone,
extending beyond the riverside mounds, charac-
teristic of the Ferlo region.
In the 1970s, Poulet (1982) collected pellets in the
Ferlo region, at Fété-Olé (16°14’N, 15°06’W), but
during our survey in this area in 2002 we did not
find a single pellet. We therefore decided to explore
farther south along a line stretching between the
towns of Louga and Linguère. In August 2003, we
sampled six different sites: Louga (15°36’N;
16°12’W), Thiamene (15°28’N, 15°52’W), Mouk-
Mouk (15°26’N, 15°46’W), Boulal (15°23’N,
15°39’W), Dahra-Djolof (15°20’N, 15°28’W) and
Nguith (15°24’N, 15°09’W). The total annual rainfall
recorded at Linguèr in 2003 was 414.5 mm. This is
far lower than that recorded before the 1970s: aver-
age annual rainfall between 1951 and 1969 was
524 mm (S.D. = 117; min/max = 33/679) and de-
creased to 367 mm (S.D. = 93; min/max = 191/587)
between 1970 and 2000.
Pellets were collected at owls’ nesting sites,
located in abandoned buildings (former railroad
stations, old houses, barns, etc.), under big trees
and particularly on the landings of water towers.
Identification of prey
The prey species in the pellets were identified
under a binocular magnifier, based either on the
morphology or morphometrics of whole or frag-
mented skulls, mandibles and teeth, by comparison
with a reference collection (IRD Sénégal) and also
with drawings in Rosevear (1969). Skulls and man-
dibles are deposited at the CBGP Laboratory in
Dakar. In the genus Mastomys, two sibling species
are sympatric in the Richard-Toll area, and cannot
be distinguished morphologically or morpho-
metrically, but have different habitat requirements
(Duplantier et al. 1988): M. erythroleucus inhabits dry
areas and villages, whereas M. huberti is restricted
to humid areas (rice fields and swampy areas
along the river). Identification of these species was
thus based on the type of habitat where the pellets
were collected, Conversely, the two species of the
genus Taterillus are sympatric and syntopic: in this
case there is no means to distinguish the two
species morphometrically or ecologically, so iden-
tification was not possible beyond the generic
level. For the genus Gerbillus, two species of very
different sizes were known in the early 1990s
(Duplantier et al. 1991), but a third species of inter-
mediate size appeared at the end of the 1990s (Bâ
2002). We discovered no qualitative morphological
137
trait allowing us to differentiate the skulls, but
adults can be distinguished by their size; we there-
fore used mandible size distributions to determine
whether two or three species were present at a
given location and at a given date. As far as shrews
are concerned, species determination is difficult on
morphological criteria, since they all belong to the
genus Crocidura. We therefore took various mea-
surements using a slide calliper: total length of
skull, largest width of mandible, interorbital con-
striction, greatest width of skull cap, upper and
lower rows of teeth and mandible height at the
coronoid process. According to identification of
individuals collected by trapping and owl pellet
analysis during the eighties (Hutterer, Granjon &
Duplantier, unpubl. data), at least five species
occur in this area: C. foxi, C. fuscomurina, C. lusi-
tania, C. nanilla and C. viaria. But as specific identi-
fications of shrews collected recently (2000–2003)
have not yet been confirmed, it is not possible to
evaluate the changes in shrew communities.
Data analysis
We determined the total number of prey per
species, per site and per date, but not the number
of prey per pellet since many pellets were already
decomposed at some localities. To detect changes
in the small mammal communities over time, we
compared our results from the Richard-Toll site to
data obtained between April 1989 and March 1990
from owl pellets collected in the Djoudj National
Parc (70 km west of Richard-Toll in the Senegal
delta, Bâ et al. 2000). We also compared our results
with data obtained from trappings between July
1990 and July 1993 by Duplantier & Sène (2000), as
well as to data from Bâ (2002) obtained between
July 1998 and May 2001 from trappings and
captures by hand around the town of Richard-Toll.
Along the Louga-Linguère line, we compared
our results with data obtained by trapping and
from pellets collected between 1970 and 1977 by
Poulet (1982) in Fété-Olé (northern Ferlo) and also
with unpublished data of Bâ & Duplantier ob-
tained by trapping and hand-capture in 1993 and
2003 around Barkedji (15°16’N, 14°51’W) located
about 30 km east of Linguère.
RESULTS
We analysed a total of 2526 whole pellets, 1808 of
which were from Richard-Toll (Diéry and Walo
zones) and 718 from the Louga-Linguère line
(Ferlo region), as well as decomposed pellets.
138 African Zoology Vol. 43, No. 2, October 2008

Table 1. Contents of barn owl (Tyto alba) pellets at the Richard-Toll airfield (Diéry – dry areas) between 1989 and
2002: percentages per animal group and total number of individuals.

1989 1991 1992 1993 1998 2001 2002

Rodents (%) 68 97 97 96 89 95 97
Shrews (%) 5 1 2 2 6 4 2
Bats (%) 0 0.1 0 0 0 0.4 0.5
Birds (%) 27 2 1 2 3 0.4 0.5
Amphibians (%) 0 0 0 0 1 0 0
Total number of prey items 52 767 1009 146 1158 241 822

Richard-Toll from 1989 to 2002
In the Diéry zone (Table 1), among 4195 identi-
fied prey of barn owls, small mammals and partic-
ularly rodents comprised the major part of the
predators’ food. Over the seven years of field-
work, rodents consistently represented the most
abundant prey, while shrews ranked second.
Among the latter, Crocidura lusitania (37%) was the
most abundant species and the only one always
represented, while C. viaria (33%) and C. nanilla
(21%) were absent in 1991 and 1993, respectively.
Bats were present only in 1991, 2001 and 2002.
Birds made up over one quarter of the predator ’s
diet in 1989, but presented only very low frequencies
as of 1991. Amphibians were present in the owls’
diet only in 1998. Only 9% of specimens could not
be reliably identified.
Seven rodent genera were identified (Table 2),
six of which belong to the Muridae family and one
to the Dipodidae family. The owls’ diet is almost
entirely made up of Muridae: the sub-family
Gerbillinae (85%) is largely dominant over Murinae
(15%). The genus Gerbillus (77% of pellets on
average) represents the main prey, Desmodilliscus
braueri and the genus Taterillus rank second with
low average frequencies (5.3% and 2.2%, respec-
tively). Between 1989 and 2002 the percentage of
Taterillus decreased from 15% to 2.4%, while that
of D. braueri increased from 3% to 12%. Among the
Murinae, Mastomys erythroleucus and Arvicanthis
niloticus, although never abundant in the owls’
diets, were present every year. The rarest species
was Jaculus jaculus (Dipodidae) represented by
only one specimen. The frequencies of Mastomys
sp., Arvicanthis niloticus and Mus (Nannomys) sp.
markedly declined over time, while the frequency
of Desmodilliscus braueri increased by over 10%
between 1993 and 2002.
To examine how many species of the genus
Gerbillus were present at Richard-Toll, we measured
the length of 300 mandibles, i.e. 100 for each of the
years 1991, 2001 and 2002 (Fig. 2). The measurements
taken in 1991 followed a bimodal distribution,
representing distinct small and large species. The
200 measurements taken in 2001 and 2002 were
distributed into three peaks, with the appearance
of a medium-sized species. These histograms also
show that the new species is now as abundant as
the two others at Richard-Toll (36% of gerbils).
In the rice fields (Walo wetlands) of Colonat-
Richard-Toll (Table 3), small mammals (86% of
total) were represented mainly by rodents, which
were the most abundant animals in every sampling
year, with frequencies reaching 100% at times

Table 2. Numbers of individuals per rodent species in barn owls pellets at the Richard-Toll airfield (Diéry – dry areas)
between 1989 and 2002.

1989 1991 1992 1993 1998 2001 2002

Mastomys spp. 10 4 123 107 78 57 98

Arvicanthis niloticus 5 2 22 12 11 7 19
Mus (Nannomys) spp. 2 0 26 5 1 1 1
Taterillus spp. 4 3 7 0 13 5 14
Gerbillus spp. 12 734 759 13 768 141 561
Desmodilliscus braueri 1 4 11 3 107 15 93
Jaculus jaculus 0 0 1 0 0 0 0
Total 34 747 949 140 978 226 786
 Thiam et al.: Impacts of climate changes on Sahel small mammal communities 139

Fig. 2. Appearance of a third species of gerbil, evidenced by mandible lengths for Gerbillus spp. collected in owl
pellets: bimodal histogram, at Richard-Toll in 1991 (A); trimodal histograms at Richard-Toll in April 2001 (B) and Octo-
ber 2002 (C), and at Nguith in 2003 (D). All graphs at same scale.

(1993), and by shrews with frequencies varying
between 7% and 9%. C. viaria was the most abun-
dant shrew species (75%) and was present every
year, while C. nanilla (13%) and C. lusitania (8%)
were found only in 1989. Birds were present only
in 1989 (44%) and in 2002 (14%), while bats and
amphibians were never found at this location.
The subfamily Murinae (four species) is clearly
dominant over the Gerbillinae (one species) in this
habitat (Table 4). Mastomys huberti and Arvicanthis
niloticus were the most abundant prey species.
Based solely on skull morphology, M. huberti
cannot be distinguished from M. erythroleucus, but
the bimonthly captures carried out in this habitat
140 African Zoology Vol. 43, No. 2, October 2008

Table 3. Contents of barn owl pellets in Colonat- Table 4. Numbers and relative frequencies of different
Richard-Toll (Walo zone wetlands and rice fields) rodent species in barn owl pellets collected at Colonat-
between 1989 and 2002: percentages per animal group Richard-Toll (wetlands and rice fields) from 1989 to
and total number of prey. 2002.

1989 1993 2001 2002 1989 1993 2001 2002

Rodents (%) 47 100 93 78 Mastomys spp. 25 91 50 45

Shrews (%) 9 0 7 8 Arvicanthis niloticus 7 30 30 38
Birds (%) 44 0 0 14 Mus musculus 0 0 0 2
Mus (Nannomys) spp. 0 1 0 0
Total numbers 75 122 87 110 Gerbillus spp. 2 0 1 1
Undetermined rodents 1 0 0 0
Total rodents 35 122 81 86

Table 5. Contents of barn owl pellets in six villages along the Louga-Linguère road in 2003: percentages per animal
group and total number of prey.

Nguith Dahra-Djolof Boulal Mouk-Mouk Thiaméne Louga

Rodents (%) 92 90 87 99 98 100

Shrews (%) 1.3 2.3 6 0 0 0
Bats (%) 0.4 0 1 0 1 0
Birds (%) 6 6 7 1.2 1 0
Amphibians (%) 0.2 1 0 0.3 0 0
Total number of prey 535 297 391 339 191 61

during a period of three years as part of a study on
schistosomiasis (over 1500 individuals) showed
that only M. huberti is present in these wetlands
(Duplantier & Sène 2000).
Mice, Mus (Nannomys) sp. and Mus musculus
were present in the owls’ diets only in 1993 and
2002, with respective frequencies of 1% and 2%.
The genus Gerbillus (Gerbillinae) made up only 2%
of the total.
The Ferlo in 2003: Louga–Linguère line
Among the 1814 identified prey of barn owls
(Table 5), rodents were the most abundant (94%).
They were present at all sites with frequencies
above 87% in all cases. Birds represented only
1–7% of prey, and were absent from one site.
Shrews were present at three sites: C. lusitania was
the most abundant (71%), followed by C. nanilla
(23%) and C. viaria (6%).
As far as rodents are concerned (Table 6), six
genera belonging to the Muridae were identified,
and the vast majority of prey items (96%) were
Gerbillinae. The genus Gerbillus (88%) was the
most abundant at all sites. Based on measure-
ments of 120 mandibles of Gerbillus collected at
Nguith (Fig. 2D), there were three species of differ-

Table 6. Numbers of individuals per rodent species in barn owl pellets in six villages along the Louga–Linguère road in
August 2003.

Nguith Dahra-djolof Boulal Mouk-Mouk Thiaméne Louga Tota l

per species

Arvicanthis niloticus 0 2 12 0 0 0 14
Mus musculus 1 2 0 0 0 0 3
Mus (Nannomys) spp. 1 15 31 1 0 1 49
Taterillus spp. 0 31 64 1 4 16 116
Gerbillus spp. 488 221 224 332 184 44 1493
Desmodilliscus braueri 8 0 8 3 2 0 21
Total rodents 498 271 339 337 190 61 1696
 Thiam et al.: Impacts of climate changes on Sahel small mammal communities
ent sizes present. The new species of intermediate
size seems to be less abundant at the moment than
in northern Richard-Toll (only 28% of gerbils). The
genus Taterillus (7%) was the next most frequent
prey item, although it was absent from one site.
Desmodilliscus braueri (1%) appeared in only four
sites and in very low numbers. Arvicanthis
niloticus, Mus musculus and Mus (Nannomys) sp.
were the only representatives of the Murinae.
Trapping results in the Ferlo area
In Barkédji, a locality east of Linguère (c. 30 km),
captures carried out by Bâ and Duplantier in 1993
(unpubl. data) included a total of 48 individuals:
Taterillus sp. represented 62.5%, Arvicanthis
niloticus 33% and Mastomys erythroleucus 4%. Two
Desmodilliscus braueri and one Taterillus were also
captured during nightly trips. In 2003, captures
included a total of 60 individuals, comprising 80%
A. niloticus, 11.7% Taterillus, 3.3% Gerbillus and 5%
Mastomys erythroleucus. During nightly trips three
D. braueri and one Gerbillus henleyi were captured
by hand. Here again, a decrease in Taterillus (and
the appearance of Gerbillus) is evidenced over the
past decade.
DISCUSSION
Our results corroborate the existing data on barn
owl pellets for this sub-region in showing that
small mammals, and particularly rodents, make
up the main part of their diet: they represent 98%
of the prey in Mauritania (Poulet 1974), about 90%
in Mali (Wilson 1987) and 75% in Nigeria (Demeter
1978; Lekunze et al. 2001). In Senegal, Poulet (1982)
described barn owl diets in detail for the period
1971–1977 in the Ferlo region, and rodents repre-
sented about 50%. In Djoudj National Parc
(Senegal Delta), Bâ et al. (2000) showed that
rodents made up nearly 88% of the owl’s food.
In wetlands along the Senegal River, M. huberti is
the dominant species, as well as in Mali. In the
Niono region in Mali (rice fields and market
gardens along the Sahel Canal), Wilson (1987) also
showed that Mastomys represented over 80% of
the rodent prey. Likewise, at the Batamani site
(lower Niger Delta, Mali), Granjon and Traoré
(2007) showed that Mastomys huberti represented
76.5% of micro-mammal prey species, followed by
the Crocidura (11.1%). A second species of
Murinae, A. niloticus, was present yet rare in owl
pellets due to its mostly diurnal habits (Duplantier
& Granjon 1991).
Measurements of Gerbillus mandibles (Table 2)
141
from the Richard-Toll and Louga-Linguere areas
confirmed that three species are indeed present:
G. henleyi (small) and G. tarabuli (large), known
from the Richard-Toll zone since 1989 (Duplantier
et al. 1991); and G. nigeriae, of intermediate size,
first captured in 1999 (Bâ et al. 2006). In 1993, the
genus Mastomys probably experienced an out-
break, since it represented over 75% of rodent
prey. The species was most likely M. erythroleucus,
since M. huberti is strictly limited to humid areas.
Arvicanthis niloticus was rare in owl pellets, proba-
bly owing to its diurnal habits, but trapping
between 1998 and 2001 in this part of Diéry
showed that A. niloticus was indeed present,
accounting for 48% of captures, followed by
Gerbillus (39%) (Bâ 2002). Desmodillicus braueri
(Gerbillinae), a typically Sahelian species
(Duplantier et al. 1997), ranked after Mastomys, yet
in contrast to the latter, its abundance increased
between 1993 and 2002. Not a single D. braueri was
found during captures by Bâ (2002), which is not
surprising since this species is known to avoid
traps (Poulet 1984). It was, however, captured by
hand at night (Bâ 2002). The genus Taterillus is
found currently far less frequently than during the
1970s (Poulet 1982).
In the 1970s in the same area, Murinae were very
infrequent; not a single Mastomys was identified,
and A. niloticus represented only 1% of a total of
1696 identified rodents (Poulet (1982), while at
Fété-Olé (see location on Fig. 1) shrews represented
over 70% of prey between 1970 and 1971, followed
by Taterillus sp. (27%). Such large proportions of
shrews are exceptional and difficult to explain for
such a dry environment. Following the 1972
drought, i.e. between 1974 and 1976 in this location,
Poulet (1982) showed that Taterillus made up the
main part of the raptor ’s food, with 65% of prey,
followed by Crocidura (29%). In 2003, the presence
of Taterillus in owl pellets shrunk to 6.7% of the
total prey. Trapping results for Barkedji confirm
the pellet analysis: Taterillus was decreasing in
number from 1993 to 2003, while the murine A.
niloticus increased markedly. Not a single Gerbillus
was found in the pellets, neither between 1970 and
1971, nor between 1974 and 1976 (Poulet 1982). Yet
in 2003, they were the main prey of barn owls in
northern Senegal, representing 88% of the total.
The two gerbil species (G. tarabuli and G. henleyi),
which arrived in northern Senegal in the late 1980s
(Duplantier et al. 1991), are now abundant in the
Ferlo, and have recently even been captured as far
south as Bandia (80 km south of Dakar) by Bâ
142 African Zoology Vol. 43, No. 2, October 2008
(2002). Gerbillus nigeriae, reported for the first time
in Senegal in Richard-Toll in 1999 (Bâ 2002), is now
present in the Ferlo and is as abundant as the two
other species.
Changes in geographic distribution of certain
species could very well be the consequence of
climatic changes that occurred in this region: the
isohyets clearly shifted southwards between the
periods 1947–1969 and 1970-1992 over the entire
country (see Fig. 1: from Trape et al. 1996). From
the north, we have observed the appearance of
four new species in the country, and on the oppo-
site, other species such as Desmodilliscus braueri
(Duplantier et al. 1997) and Taterillus gracilis
(Thiam et al. work in progress), are moving south-
wards, from the northern part of the country to
the Gambia.
In the arid zones of northern Senegal, the increase
in numbers of rodents adapted to arid conditions
(gerbils and jerboas), the appearance of a new
species in the region (Gerbillus nigeriae, Bâ et al.
2006) and the decrease in abundance of murines
and shrews, which are more adapted to humid
climates and habitats, show that this area is
becoming increasingly arid. Gerbils and jerboas
are not commensal species benefiting from human
movements to colonize new territories, but feral
species whose ranges appear to be increasing
naturally (without human facilitation). Given this,
the extent of range extension is remarkable:
250 km in less than 20 years for G. tarabuli, and
150 km in about 10 years for G. nigeriae. The
distribution area of gerbils now covers about half
of the country. This wide distribution attained by
these species in a so short a time shows that the
desertification process that began in this area in
the early 1970s (Handschumacher et al. 1992) is
continuing as a long-term process. Gerbils are
clearly good biological indicators of the marked
ecological modifications in the Sahel recently.
Future research should investigate why the
range of Gerbillus is extending and replacing that
of Taterillus. We intend to explore several possibili-
ties, namely that Gerbillus may have introduced
and spread pathogens lethal to Taterillus; that the
endemic species are ill-adapted to the new climatic
conditions; or perhaps the invading species
outcompete the endemic gerbils. To determine the
origin of these invading populations, we shall
compare, by karyology and molecular biology, the
gerbils from Senegal and those from neighbouring
countries (Mali, Mauritania) as potential source
populations. Preliminary karyological analysis (Ba
et al. 2006) favours the hypothesis of a Mauritanian
origin for G. nigeriae.
AKNOWLEDGEMENTS
We thank O. Niang for help in collecting pellets,
Y. Niang for help in dissecting pellets, L. Granjon
for comments on an earlier version of this manu-
script and E. Taube for translation.
REFERENCES
BÂ, K. 2002. Systématique, écologie et dynamique de popula-
tions de petits rongeurs potentiellement réservoirs ou hôtes
de virus au Sénégal. Mémoire de diplôme EPHE,
Montpellier.
BÂ, K., GRANJON, L., HUTTERER, R. & DUPLANTIER,
J.M. 2000. Les micromammifères du Djoudj (delta du
Sénégal) par l’analyse du régime alimentaire de la
chouette effraie, Tyto alba. Bonner Zoologische Beitrage
49: 31–38.
BÂ, K., THIAM, M., DOBIGNY, G., GRANJON, L.,
MANÉ, Y., VOLOBOUEV, V. & DUPLANTIER, J.M.
(2006). Hypotheses on the origin of the invasion of
Senegal by Gerbillus nigeriae based on chromosomal
data. Mammalia 70(3/4): 303–305.
DEMETER, E. 1978. Food of a barn owl Tyto alba in
Nigeria. Bulletin of the Nigerian Ornithological Society
41(4): 9–13.
DRAPEAU, P., LEDUC, A. & BERGERON, Y. 2003. Bird
communities in old lichen-black spruce stands in the
clay belt: Problems and solutions regarding forest
management. Forestry Chronicle 79(3): 531–540
DUPLANTIER, J.M. & GRANJON, L. 1988. Occupation
et utilisation de l’espace par des populations du
genre Mastomys au Sénégal: étude à trois niveaux de
perception. Sciences et. Techniques de l’Animal de
Laboratoire 13(2): 129–134.
DUPLANTIER, J.M. & GRANJON, L. (1991). Rythmes
d’activité chez six espèces de Muridés du Sénégal
appartenant aux genres Mastomys, Avicanthis, Myo-
mys et Dasymys. Mammalia 54(2): 173–182.
DUPLANTIER, J.M. & SÉNE, M. 2000. Rodents as
reservoir hosts in the transmission of Schistosoma
mansoni in Richard-Toll, Senegal, West Africa. Journal
of Helminthology 74: 129–135.
DUPLANTIER, J.M., GRANJON, L. & BÂ, K. 1991.
Découverte de trois espèces de rongeurs nouvelles
pour le Sénégal: un indicateur supplémentaire de la
désertification dans le nord du pays. Mammalia 55(2):
313–315.
DUPLANTIER, J.M., GRANJON, L. & BÂ, K. 1997.
Répartition biogéographique des petits rongeurs au
Sénégal. Journal of African Zoology 111(1): 17–26.
GRANJON, L. & TRAORÉ, M. 2007. Prey selection by
barn owls in relation to small-mammal community
and population structure in a Sahelian agro-eco-
system. Journal of Tropical Ecology 23: 199–208.
GOETZE, D., HORSCH, B. & POREMBSKI, S. 2006.
Dynamics of forest-savanna mosaics in north-eastern
Ivory Coast from 1954 to 2002. Journal of Biogeography
33(4): 653–664.
HANDSCHUMACHER, P., HERVÉ, J.P. & HÉBRARD, G.
1992. Des aménagements hydro-agricoles dans la
 Thiam et al.: Impacts of climate changes on Sahel small mammal communities
vallée du fleuve Sénégal ou le risque des maladies
hydriques en milieu sahélien. Sécheresse 3(4): 219–
226.mm
HEIM DE BALSAC, H. 1965. Quelques enseignements
d’ordre faunistique tirés de l’étude du régime
alimentaire de Tyto alba dans l’Ouest de l’Afrique.
Alauda 33: 309–322.
HODKINSON, I.D. 2005. Terrestrial insect along eleva-
tion gradients: species and community responses to
altitude. Biological Review 80(3): 489–513.
HULME, M. 1992. Rainfall changes in Africa: 1931–1960
to 1961–1990. International Journal of Climatology 12:
685–699.
HULME, M., BARROW, E.M., ARNELL, N., HARRISON,
P.A., DOWNING, T.E. & JOHNS, T.C. 1999. Relative
impacts of human-induced climate change and natu-
ral climate variability. Nature 397: 588–591.
HULME M., DOHERTY, R., NGARA, T., NEW, M. &
LISTER, D. 2001. African climate change: 1900–2100.
Climate Research 17: 145–168
LEKUNZE, J.M., EZCALOR A.U. & AKEN’OVA, T. 2001.
Prey groups in the pellets of the barn owl Tyto alba
(scopoli) in the Nigerian savanna. African Journal of
Ecology 39: 38–44.
LEROUX, M. 1983. Climat. In: Atlas du Sénégal, (ed.) P.
Pélissier, pp. 12–17. Editions Jeune Afrique, Paris.
MALEQUE, M.A., ISHII, H.T. & MAETO, K. 2006. The use
of arthropods as indicators of ecosystem integrity
in forest management. Journal of Forestry 104(3):
113–117.
MORON, V. 1994. Guinean and Sahelian rainfall anom-
aly indices at annual and monthly scales (1933–1990).
International Journal of Climatology 14: 325–341.
NICHOLSON, S.E. 1978. Climatic variations in the Sahel
and other African regions during the past five centu-
ries. Journal of Arid Environments 1: 3–24.
POPOV, G.B. 1996. Quelques effets de la sécheresse
sahélienne sur la dynamique des populations acri-
diennes. Sécheresse 7: 91–97.
POULET, A.R. 1972a. Caractéristiques spatiales de
Taterillus pygargus dans le Sahel sénégalais. Mammalia
36(4): 579–606.
143
POULET, A.R. 1972b. Recherches écologiques sur une
savanne sahélienne du Ferlo septentrional, Sénégal:
Mammifères. Terre & Vie 26(3): 440–472.
POULET, A.R. 1974. Rongeurs et insectivores dans les
pelotes d’effraie en Mauritanie. Mammalia 38: 10–11.
POULET, A.R. 1982. Pullulation de rongeurs dans le Sahel.
Mécanismes et déterminisme du cycle d’abondance
de Taterillus pygargus et d’Arvicanthis niloticus (Ron-
geurs, Gerbillidés et Muridés) dans le Sahel du Séné-
gal de 1975 à 1977. Ph.D. thesis, Université Paris VI.
POULET, A.R. 1984. Quelques observations sur la
biologie de Desmodilliscus braueri Wettstein (Roden-
tia, Gerbillidae) dans le Sahel du Sénégal. Mammalia
48: 59–64.
ROMBKE, J., JANSCH, S. & DIDDEN, W. 2005. The use of
earthworms in ecological soil classification and
assessment concepts. Ecotoxicological Environment
Safety 62(2): 249–65.
ROSEVEAR, D.R. 1969. Rodents of West Africa. Trustees of
the British Museum (Natural History), London.
SAINT-GIRONS, M.C. & SPITZ, F. 1966. A propos de
l’étude des micromammiferes par l’analyse des
pelotes de rapaces: intérêt et limites de la méthode.
Terre & Vie 4: 3–17.
TAYLOR, I. 1994. Barn Owls. Predator–prey relationship and
conservation. Cambridge University Press, Cambridge.
TRAPE, J.F., GODELUCK, B., DIATTA, G., ROGIER, C.,
L E GR O S , F. , A L B E R GE L , J . , P E P I N , Y. &
DUPLANTIER, J.M. 1996. The spread of tick-borne
borreliosis in West Africa and its relationship to
Sub-Saharan drought. American Journal Tropical
Medecine Hygiene 54(3): 289–293.
WHITFIELD, A.K. & ELLIOTT, A. 2002. Fishes as indica-
tors of environmental and ecological changes within
estuaries: a review of progress and some suggestions
for the future. Journal of Fish Biology 61(17): 229–250.
WILSON, R.T. 1987. Le régime alimentaire de la chouette
effraie, Tyto alba au Mali central. L’oiseau et R.F.O. 57(3):
195–200.
YOM-TOV, Y. & WOOL, D.N. 1997. Do the contents of
barn owl pellets accurately represent the proportion
of prey species in the field? Condor 99: 972–976.
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