On the western fringe of baboon distribution: mitochondrial D-loop
diversity of Guinea Baboons (Papio papio Desmarest, 1820) (Primates:
Cercopithecidae) in Coastal Guinea-Bissau, western Africa
Maria J. Ferreira da Silva *1,2,
Catarina Casanova 2,3,4,5 & Raquel Godinho 1
1 CIBIO/InBio, Centro de Investigação em Biodiversidade e
Recursos Genéticos, Universidade do Porto, Campus Agrário de Vairão, 4485-661
Vairão, Portugal.
2 APP, Associação Portuguesa de Primatologia, Av. Berna, 26-C, 1069-061 Lisboa, Portugal
3 CAAP, Centro de
Administração e Políticas Públicas, Instituto Superior de Ciências Políticas e
Sociais da Universidade Técnica de Lisboa, Pólo Universitário do Alto da Ajuda,
R. Almerindo Lessa, 1300-663 Lisboa, Portugal
4 Department of
Anthropology, Instituto Superior de Ciências Sociais e Políticas (ISCSP) da
Universidade Técnica de Lisboa, Pólo Universitário do Alto da Ajuda, R.
Almerindo Lessa, 1300-663 Lisboa, Portugal
5 CESAM, Centre for Environmental and Marine
Studies, Lisbon Group, Portugal
1 ferreiradasilvamj@cf.ac.uk (*
corresponding author), 2 ccasanova@iscsp.utl.pt, 3 rgodinho@cibio.up.pt
doi: http://dx.doi.org/10.11609/JoTT.o3216.4441-50| ZooBank: urn:lsid:zoobank.org:pub:389730A6-1E26-4486-A07C-CF62279AD32A
Editor: Dietmar Zinner, German Primate
Center, Göttingen, Germany. Date
of publication: 26 June 2013 (online & print)
Manuscript details: Ms #
o3216 | Received 24 May 2012 | Final received 30 May 2013 | Finally accepted 01
June 2013
Citation: Ferreira da Silva, M.J., C. Casanova & R. Godinho(2013). On the western fringe of
baboon distribution: mitochondrial D-loop diversity of Guinea Baboons (Papio
papio Desmarest, 1820) (Primates: Cercopithecidae) in Coastal
Guinea-Bissau, western Africa. Journal of Threatened Taxa 5(10): 4441–4450; http://dx.doi.org/10.11609/JoTT.o3216.4441-50
Copyright: © Ferreira da Silva et al. 2013. Creative Commons
Attribution 3.0 Unported License. JoTT allows unrestricted use of this article
in any medium, reproduction and distribution by providing adequate credit to
the authors and the source of publication.
Funding: Funding was provided by CIBIO/InBio - Centro de
Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto and
by two private companies - Barbosa & Almeida Glass I - Serviços de Gestão e
Investimentos, SA. and Carosi - Cápsulas e Rolhas Sintéticas Lda. R. Godinho
work was funded by Portuguese Foundation for Science and Technology (FCT) under
a post-doc grant (SFRH/BPD/36021/2007).
Competing Interest: None.
Acknowledgements: We gratefully acknowledge S. Camará, C. Sousa and A. Barata for
assistance in fieldwork. We thank Direcção Geral das Florestas e Faunado Governo da República da Guiné-Bissau, IBAP (Instituto da Biodiversidade e
Áreas Protegidas) and AD (Acção para o Desenvolvimento, NGO) for
helping in the logistics of the study. We acknowledge ICNB - Divisão de
Aplicações de Convenções/CITES (Biodiversity and Nature Conservation
Portuguese Institute/ CITES convention application division) for sample
transport permits to Portugal. Photo courtesy of A. Barata. We also thank G.
Dantas and CTM staff for lab assistance. We acknowledge A. Beja-Pereira, I.
Russo, J. Cable, J. Chen, N. Ferrand and S.J.E. Baird for helpful discussions
and manuscript comments, to R. Lopes for figures design and to D. Zinner and
two anonymous reviewers whose comments greatly improved this manuscript.
Molecular work was supported by CIBIO/InBio. Fieldwork was sponsored by Barbosa
& Almeida Glass I - Serviços de Gestão e Investimentos, SA. and Carosi -
Cápsulas e Rolhas Sintéticas Lda. APP (Portuguese Primatological Association) and DARI Project assured
fieldwork logistics. R. Godinho worked under a FCT post-doc grant
(SFRH/BPD/36021/2007).
Author Contribution: Ferreira da Silva was responsible for the field
and laboratorial work, data analysis and writing of the manuscript. All
co-authors were Ferreira da Silva master supervisors and contributed to the
planning of the project and provided the important suggestions during the
manuscript preparation.
Author Details: Maria
J. Ferreira da Silva was a master student in CIBIO (Research
Centre in Biodiversity and Genetic Resources, Porto University) and has been researching
Guinea baboons in Guinea-Bissau for the last seven years. She finished her PhD
degree at School of Biosciences, Cardiff University, UK in 2012 and is
currently a post-doc researcher in Cardiff University and in CIBIO.
Catarina Casanova is an associated professor in the Department of Anthropology at the
School of Social and Political Sciences (Technical University of Lisbon,
Lisbon, Portugal). She completed her DPhil degree in the University of
Cambridge, UK. She has being working in Guinea-Bissau for the last eight years
and is specialized in biological anthropology (Primatology), ethnobiology and
primate conservation.
Raquel Godinho is post-doc researcher in
CIBIO. She completed her PhD degree in Faculty of Sciences in Lisbon
University, Portugal. She works in conservation genetics for the last six years
and has in-depth experience in analyzing non-invasive biological samples of
various mammal species.
Abstract: Like many primate species in West Africa, habitat loss and intensive
hunting are threatening the poorly studied Guinea Baboon (Papio papio).
These factors contributed to a significant population contraction during the
last 30 years. Our study presents
genetic diversity estimates for the Guinea Baboon based on a 391 base pair
fragment of the mitochondrial DNA D-loop hypervariable region I. We used non-invasively collected genetic
samples from two locations in Guinea-Bissau: Cufada Lagoons Natural Park and
Cantanhez Forest National Park. Although most sampling was opportunistic, we observed and collected
samples from two dames (social units). Among the 25 sequences obtained, we
found seven closely related mtDNA haplotypes and one highly different
haplotype. The presence of this
divergent haplotype suggests a contact area between genetically differentiated
populations in Cufada Lagoons Natural Park, or dispersal of individuals. The samples gathered from both regions
share two of the most common haplotypes in different frequencies, but also exhibit
unique haplotypes. No significant
genetic differentiation was found between social units from both regions,
possibly due to common ancestral origin or frequent dispersal between sampling
locations. The presence of
different maternal lineages in the same social unit and a higher percentage of variation
within social units suggest historical female-biased dispersal for
Guinea-Bissau Baboons. We further
compared mitochondrial genetic diversity of Guinea and Hamadryas Baboons. We found lower haplotype, nucleotide
and theta diversity for Guinea Baboons, which points to different demographic
histories of these species. This
work supports the need for additional genetic studies within the full Guinea
Baboon range.
Keywords: Baboon, dispersal, habitat loss,
hunting, Mitochondrial DNA, primates.
For figures, images, tables -- click here
Introduction
The free-ranging populations of Guinea
Baboons (also called Red Baboons, Papio papio Desmarest, 1820) occupy a
small distribution area in western Africa (Mauritania, Senegal, Gambia, Mali,
Guinea-Bissau, Republic of Guinea and Sierra Leone) when compared with other
baboon species (Oates et al. 2008). This poorly studied primate exhibits great ecological plasticity
(Galat-Luong et al. 2006; Patzelt et al. 2011) and inhabits a variety of
habitats throughout its distribution (Galat-Luong et al. 2006).
Guinea Baboons have been listed as Near
Threatened and little is known about its populations (IUCN 2013 assessed by Oates
et al. 2008) (Image 1). It is
acknowledged that certain local or regional populations deserve conservation
measures due to their rapid decline in numbers. In several areas in western Africa only
fragmented populations persist (Galat et al. 1999–2000). Excessive habitat degradation by
agricultural practices (e.g. slash and burn technique), hunting and persecution
by farmers, international trade of juveniles and bushmeat consumption (Wolfheim
1983; Starin 1989; Galat et al. 1999–2000; Oates et al. 2008; Minhós et
al. under review) contributed to substantial range contraction during the last
30 years (Oates et al. 2008).
In Guinea-Bissau, wild populations of
Guinea Baboons seem to be decreasing at a fast pace (Gippoliti & Dell’Omo
2003; Casanova & Sousa 2007; Cá 2008; Costa 2010). In the 1980s it was still possible to
regularly observe troops of Guinea Baboons in most regions of the country and
their range extended into the outskirts of Bissau, the capital city. After this date, the baboon conservation
status in Guinea-Bissau degraded dramatically. According to the perceptions of local
people (Costa 2013) and hunters (Cá 2008; Amador in press), baboons became rare
and live in smaller groups or have even disappeared from some parts of the
country where they were common 30 years ago. This decrease could be correlated with:
(i) changes in the landscape due to extensive cashew tree (Anacardium
occidentalis) plantation, which now occupies more than 70% of the whole
arable land (Barry et al. 2007); (ii) intensive hunting by military groups that
have consumed baboon meat in return for salary (Casanova & Sousa 2007);
(iii) bushmeat markets and restaurants that flourished in the capital and in
some other smaller cities throughout the country, where baboon meat is sold and
consumed along with other primate species (Casanova & Ferreira da Silva
pers. obs. 2006–2010; Cá 2008; Starin 2010; Minhós et al. under review);
(iv) the use of baboon skins as part of folk medicine practices (Ferreira da
Silva et al. 2009; Sá et al. 2012); and (v) pet trade of very young
individuals, which is a common practice throughout the country (Casanova &
Sousa 2007; Hockings & Sousa 2011; Ferreira da Silva, pers. obs.
2006–2010). Baboons are
believed to be relatively common in the southern part of the country (although
patchily distributed) and absent from the northwest (Gippoliti & Dell’Omo
2003; Casanova & Sousa 2007; Oates et al. 2008). However, little is known about the
conservation status of these persisting populations and the impact of
anthropogenic related habitat modifications and hunting for pet and bushmeat
trade.
One possible consequence of the rapid
population decline is a reduction of genetic diversity and the isolation of
sub-populations with subsequent low levels of gene flow among them, which may
decrease the ability of the species to respond to future environmental changes
(Frankham et al. 2002). The
threats affecting baboons in Guinea-Bissau justify an assessment of the genetic
diversity and degree of gene flow between persisting populations to understand
their risk of extinction (Avise 2000) and to assist their future conservation
and management plans.
This study describes the results of the
first genetic survey on baboons in Guinea-Bissau. Sampling focussed on two of the less
deforested areas (southwestern coast) within Guinea-Bissau, where baboons were
reported to be present (Gippoliti & Dell’Omo 2003; Casanova & Sousa
2007). We aimed for a description
of the genetic diversity and a preliminary assessment of the historical genetic
structure of remaining populations. Moreover, we provide a comparison of our results with those obtained for
Hamadryas Baboons (Hapke et al. 2001). This comparative analysis would help to understand the degree of genetic
diversity of the Guinea-Bissau Baboon at the genus level. Our ultimate goal was to provide
baseline data for future conservation genetic studies in Guinea-Bissau Baboons.
Methods
Sample collection and preservation
A non-invasive genetic sampling strategy
was implemented as the capture of animals for collection of biological samples
was considered unethical, logistically impracticable and a peril to the
animal’s health (Piggott & Taylor 2003). Samples were collected in the
southwestern region of the Republic of Guinea-Bissau during 25 days along in
the dry season (January and February 2006). Sampling was concentrated in two
geographically distinct locations, 60km apart: (1) administrative region of
Quinara (11034’–11051’N & 14049’–15016’W),
most frequently in Cufada Lagoons Natural Park (Parque Natural das Lagoas da
Cufada) (CLNP) and in (2) administrative region of Tombali (11020’–11005’N
& 15006’–1404’W), predominantly in Cantanhez
Forest National Park (Parque Nacional das Florestas de Cantanhez) (CNP) (Fig. 1
and Table 1 for sampling details). As no previous information was available on the location of social units
in the sampling areas, we located individuals by detecting vocalizations,
footprints and/or scats and used information from local people, guards and park
guides. The GPS position for each
sample was registered.
In 25 days of fieldwork, a total of 38
faecal samples (17 and 21 samples from CLNP and CNP, respectively) were
collected. Most of the samples were
collected opportunistically and were spatially scattered: we collected from one to three faecal
samples in nine different sampling locations (distance within regions between
16–20 km). However, we
observed and sampled two social units: Mato de Bubatchingue (MB) and Caiquene
(Table 1). Faecal samples were
preserved by desiccation using silica beads (type II from Sigma; Wasser et al.1997). After one month in the field
samples were transferred to 99% EtOH and preserved at -200C prior to
DNA extraction.
We collected two hair samples from captive
individuals during grooming sessions and a blood sample from the carcass of a
hunted animal found in the forest (Table 1). Hair samples were preserved in 99% EtOH
and the blood sample was collected using an FTA card® (Whatman International
Ltd, Kent, England).
This project complied with the protocols
approved by CIBIO, Porto University, Portugal (for sampling of biological
material and DNA extraction of blood and faecal samples) and by the School of
Social and Political Sciences, Technical University of Lisbon, Portugal (for
survey questionnaires and interviews). Permits for research and sample export were obtained from the local
authorities (IBAP, Institute for the Biodiversity and Protected Areas and DGFC,Direcção Geral das Florestas e Fauna) and research adhered to the legal
requirements of the respective countries in which research was conducted.
DNA extraction and PCR amplification
Total genomic DNA was extracted from the
outside surface of each faecal sample using guanidine-silica solution (Gerloff
1995) followed by DNA purification through microcon YM-30 columns (Millipore
Iberica S.A.U., Madrid, Spain). DNA
from hair samples was extracted using standard salting-out extraction
protocols. DNA preserved in FTA card® (Whatman International Ltd) was extracted
according to the instructions given by the manufacturer. Non-invasive samples
were processed in a laboratory dedicated to low-quality DNA samples at the
research centre CIBIO (Research Center in Biodiversity and Genetic Resources). Negative controls were always included
to monitor for cross-contamination.
A 391 base pair (bp) fragment of the
hypervariable region 1 (HVRI) of the mtDNA (D-loop) region was PCR amplified
using primers from Hapke et al. (2001). This genetic marker was selected because it: (i) is suitable to provide
an overview of the intraspecific mitochondrial genetic diversity (Wan et al.2004); (ii) allows for a higher amplification rate on non-invasive DNA samples,
when compared with nuclear DNA markers (Waits & Paetkau 2005); and (iii)
made possible a comparison of our results with those obtained for Hamadryas
Baboons (Hapke et al. 2001), one of the few available databases of D-loop
sequences for any baboon wild population.
The PCR mixture contained 0.5µM of each
primer, 4 µM of each dNTP, 3% DMSO, 1X PCR reaction buffer (with 1.9 mM MgCl2),1U of Phusion™ High-Fidelity DNA Polymerase (Finnzymes, Vantaa, Finland)
and 2µl of DNA template in a total volume of 10µl. Amplifications were performed using a
touchdown PCR protocol with an initial denaturation step at 980C for
30 sec. This was followed by 10
cycles of 10 sec at 980C, 30 sec at 580C and 15 sec at 720C,
with the annealing temperature decreasing by 0.50C each cycle, plus
30 cycles of 10 sec at 980C, 30 sec at 530C and 15 sec at
720C, with a final 5 min extension at 720C. To monitor for contaminations, all PCR
reactions included a negative and a positive PCR control, as well as the
negative control from the DNA extraction. The amplicons were sequenced using the BigDye Terminator v3.1 cycle
sequencing kit (Applied Biosystems, California, USA) on an ABI 3130xL Genetic
Analyser (16 capillary sequencer, Applied Biosystems). Samples that exhibited the same mtDNA
haplotype in each location were genotyped for three human microsatellite loci
(D10S611, D7S503 and D5S1457) known to be polymorphic in baboons (Bayes et al.2000) to test for possible repeated sampling of individuals.
We used a molecular protocol to identify
the sex of individuals (described in Ferreira da Silva 2007) and screened 22
faecal samples across five PCR replicates.
Data analysis
Sequences were manually checked for
accuracy and aligned in BIOEDIT 7.0.9 (Hall 1999). Genetic diversity indices were
calculated for the entire sequence set and for the sequences obtained from the
defined social units (MB and Caiquene) using ARLEQUIN 3.5 (Excoffier &
Lischer 2010). The number of
variable sites, the number of pairwise differences between pairs of sequences,
haplotype diversity (Hd; Nei 1987) and nucleotide diversity (π; Nei 1987) and
their respective variances were calculated for the entire sequence set.
Genetic relationships among haplotypes
were described by a median-joining network constructed using NETWORK 4.2.0.1
(Bandelt et al. 1999; www.fluxus-engineering.com). For this analysis, indels were included
and all sites were treated with equal weights. Two sequences available at GenBank were
included in the alignment for reference, one from P. hamadryas and the
other from P. papio (Accession numbers AF275457 and AF275383, respectively).
Analysis of genetic structure was
performed only with sequences from the two social units (MB and Caiquene,
N=14). This option prevented a
possible bias in the results of genetic differentiation caused by the admixture
of several social units within each geographic region. However, since some individuals of these
social groups were possibly not sampled, the number of haplotypes found in each
social group can be underestimated. We performed an Analysis of Molecular Variance (AMOVA analysis, Excoffier
et al. 1992) using ARLEQUIN 3.5 (Excoffier et al. 2010) to assess
the extent of differentiation among populations (calculating ST,CTand SC)
following a non-parametric permutation approach (10,0172 permutations).
The final dataset in our study was
compared to results obtained from Hamadryas Baboons (Hapke et al.2001). Since genetic diversity
within social groups can be influenced by the dispersal rate (Rogers 2000), we
selected pairs of Hamadryas social groups or demes located at a similar distance
to that between the two social groups sampled for Guinea Baboons (i.e., N=18
pairs of “troops”, located between 50–100 km). It is assumed that the ability to
disperse (i.e., the organism’s movement from the natal area to the breeding
area) over such a distance is the same for both species. The pairs of selected demes from Hapke
et al. (2001) were as follow: Furrus-Abdur, Furrus-Dogali,
Furrus-Debresina, Furros-Geleb, Furros-Molki, Durfo-Abdur, Durfo-Dogali,
Durfo-Molki, Durfo-Barka River, Arborobo-Abdur, Arborobo-Geleb, Arborobo-Molki,
Abdur-Dogali, Dogali-Kubkub, Debresina-Molki, Debresina-KubKub, Geleb-Kubkub
and Molki-Barka River. D-loop
sequences were downloaded from GenBank (Accession Numbers AF275383 to AF275475)
and haplotype and nucleotide diversity per site and the theta (θ) were
calculated for each selected pair using DNASP 4.10.9 (Rozas et al.2003). Finally, these values were
compared to the results from our study.
Results
DNA extraction and amplification
Extraction of DNA was attempted from all
samples collected. We successfully
amplified 31 of the collected faecal samples (82%), the two hair samples and
the blood sample for the 391 bp fragment of the mtDNA D-loop. Despite the high amplification rate achieved
for faecal samples, good quality sequences were only obtained for 26 out of the
31 amplicons. Two samples in Mato
de Bubatchingue and two samples in RM exhibited the same haplotype and the same
genotype for the three microsatellite loci. We removed from the dataset one sample
per location to prevent repetition of individuals. Using the sex identification protocol,
only 14/22 faecal samples gave consistently the same result three times. We
effectively distinguished six females and eight males among our faecal samples
of Guinea Baboons.
D-loop diversity of coastal Guinea Baboons
Sequencing of 391bp of the mtDNA D-loop region distinguished eight
haplotypes from the 25 sampled individuals (Hd=0.803±0.061). Eighteen out of 391 nucleotide sites
analyzed were polymorphic and estimated nucleotide diversity (π) was
0.0104±0.0059. Sixteen of the sites showed substitutions, with transitions
exceeding the number of transversions (15:2). Additionally, two indels were observed:
the deletion of one nucleotide in two sequences and the insertion of one
nucleotide in one sequence. The
mean number of pairwise differences between haplotypes (indels taken into
account) was 4.06±2.1.
The median-joining network built with the eight haplotypes displays two
most common haplotypes (h1 and h4), which differ by three positions and are
shared by both CLNP and CNP geographical regions but in different frequencies
(h1: 28% and 12%, and h4: 4% and 12% in CLNP and CNP, respectively) (Fig. 2). Five other haplotypes were found in lower
frequencies (h2, h5, h6, h7 and h8) and are separated by one or two mutational
steps from each other and from h1 and h4. The haplotype h8 is separated by an indel (nucleotide insertion) from
the haplotype h4. An additional
haplotype (h3) was observed in the CLNP sample. This haplotype h3 is separated from h4
by eight substitutions and one indel (nucleotide deletion), which corresponds
to a genetic divergence of 2.3% from the central haplotype h4. Both geographic regions exhibited unique
haplotypes (h2, h3 and h5 for CLNP and h6, h7 and h8 for CNP; Fig. 2).
Genetic diversity and differentiation of
social units
The two social units showed the same
haplotype diversity (Hd=0.810±0.130) and both displayed the same number of
haplotypes (n=4) (Table 1). The MB sample had slightly higher nucleotide
diversity (πMB=0.0157±0.0035) than the Caiquene sample (πCaiquene=0.00928±0.0021).
The two social groups were tested for statistically significant genetic
structure using an AMOVA analysis, which showed no phylogeographic structure of
haplotypes: 7.75% of the total variance was found between social groups whereas
most of the variance (92.25%) was present within social groups. Accordingly, the fixation index estimated
between Guinea Baboon social groups was low (Fst= 0.078) and did not differ
significantly from zero (p=0.2).
Comparison of mitochondrial variation
between Hamadryas and Guinea Baboons
The number of haplotypes found in this
study was, on average, similar to the number found in different demes of P.
hamadryas (NhHAM=6 and NhGUI=8) with similar average
sampling efforts (NseqHAM=14 and NseqGUI=14.89).
Nevertheless, all three other genetic measures were lower for Guinea Baboons
social units, as documented for haplotype diversity (average HdHAM=0.886
and HdGUI=0.846, Fig. 3i), average nucleotide diversity found per
site (πHAM=0.0356 and πGUI=0.01297, Fig. 3ii) and Theta
based on π (θπHAM=0.037±0.0053 and θπGUI=0.013,
Fig. 3iii), suggesting lower effective population size for Guinea Baboons.
Discussion
The present work estimated for the first time the mitochondrial genetic
diversity levels for wild Guinea Baboons in Guinea-Bissau and presented an
assessment of the historical genetic structure of two populations.
Genetic diversity of baboons in Guinea Bissau (HdGUI=0.803±0.061;
πGUI=0.0104±0.0059), according to Grant & Bowen (1998), may be
interpreted as a result of a large and stable population over a long period of
time or as the occurrence of a secondary contact zone between previously
differentiated lineages (Hd>0.5 and π>0.5%, Grant & Bowen 1998). In particular, the presence of a very
distinctive haplotype (h3) harboured by two individuals in a homogenous deme
(social unit Mato de Bubatchingue) suggests that the hypothesis of a contact
zone between two genetically divergent populations in CNLP (Avise et al. 1987)
best explains the observed results. Alternatively, past or contemporary dispersal from other populations can
explain the presence of h3.
Although the amplification of nuclear pseudogenes can be common in
studies of primates using mtDNA as a molecular marker (Mourier et al. 2001;
Thalmann et al. 2004), we excluded the possibility that h3 was a numt. Throughout our study and in particular
in the samples harbouring h3 (extracted from faecal samples), we did not see
ghost bands in agarose gels when testing for DNA amplification or double peaks
in chromatograms (Bensasson et al. 2001). Moreover, sequences did not show any ambiguities, either when comparing
both sequence strands or when comparing polymorphic sites between sequences in
the final alignment (Bensasson et al. 2001). We also believe that obtaining no
sequencing product for some amplicons was related to low DNA quality/quantity
and not with mutations at the flanking regions, as Hapke et al. (2001) designed
primers in highly conserved regions.
No evidence was found for significant geographic structure of
haplotypes, and we propose that baboons present in this area of Guinea-Bissau
do not represent two genetically differentiated populations. This result suggests a common origin of
CNP and CLNP populations or dispersal of individuals between the two sampling
locations. Guinea Baboons are able
to move about 40km a day (Galat-Luong pers. comm. January 2007), which almost
equals the distance between the two sampling sites in Guinea-Bissau (about
60km; see Fig. 1), and they occupy broad home ranges in Senegal (25km2,
Fickenscher 2010).
A historical female-biased dispersal pattern for the Guinea Baboon can
explain the presence of very different maternal lineages in the same social
unit (see results for CLNP social unit, Mato de Bubatchingue) and higher
percentage of variation within social units (Di Fiore 2003). This hypothesis would contrast with the
proposal of unique maternal family groups for the Senegalese population made by
Sharman (1981) based on observational data but it agrees with evidences for
male philopatry based in nuclear DNA data provided by Fickenscher (2010). Fickenscher (2010) found significantly
higher Fst values and a higher negative correlation between pairwise
relatedness and geographic distances for males when compared with females,
which imply a scenario of female-biased dispersal (Fickenscher 2010). Furthermore, in Papio male-biased
dispersal species (e.g., Gray-footed Baboons, Burrell 2008) it is common to
observe the inverse pattern of D-loop mtDNA variance found for the Guinea
Baboon (i.e., greater percentage of variance between social units than within)
and significant values of Fst between social units (e.g., Fstgra=0.757,
Burrell 2008).
Moreover, it was found (Ferreira da Silva et al. unpublished data) that
the haplotype networks of Guinea and Hamadryas Baboon social units display a
similar structure: two divergent groups were present in most Hamadryas Baboon
groups, which resembles the haplotype structure found in Guinea Baboons social
units by this study, although with a much lower degree of differentiation
between the two most frequent haplotypes (h1 and h4). Hamadryas Baboons’ display a pattern of
female-biased dispersal (Hapke et al. 2001; Hammond et al. 2006; Handley
et al. 2006) and the social organization of Guinea Baboons and Hamadryas
Baboons share some characteristics (Galat-Loung et al. 2006; Patzelt et
al. 2011). Therefore is
possible that the similarity between haplotype networks of Guinea and Hamadryas
Baboon social units is related with female-dispersal, although such proposition
needs to be further investigated.
The current work suggests lower mitochondrial genetic diversity (both
nucleotide diversity and θπ) for Guinea-Bissau Baboons when compared with
Hamadryas Baboons. Additionally,
the mean number of pairwise differences estimated for Guinea Baboons (4.06, range
12-1) is rather low when compared with those observed in Hamadryas Baboons
(12.40, range 23-1; Hapke et al. 2001). Moreover, comparing our results (πGUI=0.0104±0.0059) with
Kinda, Yellow and Gray-footed Baboons mtDNA genetic diversity estimates (πKIN=0.036,
πYEL=0.086 and πGRA =0.053, Burrell 2008), the pattern of
lower nucleotide diversity for Guinea-Bissau Baboons is still noticeable. Reasons that could explain such
differences on genetic diversity levels include: (i) a strong stochastic event
that would affect the D-loop region, as a selective sweep (Galtier et al.
2000); (ii) a past population bottleneck, which would result in a decrease in
nucleotide diversity, followed by a population expansion early enough to
generate the same number of haplotypes but not large differences among them
(Grant & Bowen 1998); or (iii) a founder effect, which could be a
consequence of a rapid colonisation of western Africa by this species (Zinner
et al. 2011) or re-colonisation events by few or genetically similar
individuals.
Disentangling among the hypotheses posed by this initial genetic survey
will be a future challenge. With
the use of more variable molecular markers (e.g., microsatellites, Wan et al.2004) it will be possible to obtain a broader idea of genetic diversity
patterns at different genome compartments and to perform studies on the
demographic history of baboons in Guinea-Bissau. Additionally, if confirmed the presence
of baboons in other areas within the country (e.g., Boé sector, in the southern
part, see Casanova & Sousa 2007), the inclusion of samples from a wider
area would allow the assessment of the degree of functional connectivity
between all persisting populations. Despite the above-mentioned limitations, our study is the first approach to estimate genetic
diversity of baboons in Guinea-Bissau and one of the very few comparing mtDNA
patterns of different species of Papio.
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