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151
Animal Biodiversity and Conservation 38.2 (2015)
© 2015 Museu de Ciències Naturals de Barcelona
ISSN: 1578–665 X
eISSN: 2014–928 X
Angelici, F. M., Mahama, A. & Rossi, L., 2015. The lion in Ghana: its historical and current status.
, 38.2: 151162.
Abstract
.— Historically, the lion population in Ghana
has been little studied and its status is poorly documented. Currently, after recent unsuccessful attempts to
nd signs of the presence of the species, many authors believe that the Ghanaian lion population is most likely
extinct. In an attempt to gather more data, since 2005 we have been carrying out lion surveys in the most
important parks and other protected areas of Ghana, mainly focusing on Mole National Park (MNP). We have
also been extensively reviewing the literature in an attempt to reconstruct the history of the presence of the
lion in the country. Although our research has not provided unequivocal evidence of the presence of the lion,
we have collected circumstantial evidence that suggests that a small lion population might still be present in
MNP and its surrounding areas.
Key words: Lion, , Ghana, Status, Mole National Park
Resumen
.— Históricamente, la población de león () en
Ghana ha sido poco estudiada y su situación actual está poco documentada. Tras los últimos intentos infruc-
tuosos de encontrar indicios de la presencia de la especie, son numerosos los autores que opinan que la
población de león en Ghana está prácticamente extinguida. En un intento por recabar más datos, desde 2005
hemos venido realizando estudios sobre esta especie en los parques más importantes de Ghana y otras zonas
protegidas del país, que se han centrado principalmente en el Parque Nacional de Mole (MNP). Asimismo,
hemos examinado los datos publicados con el n de reconstruir la historia de la presencia del león en el país.
Si bien nuestra investigación no aportó datos inequívocos, se recabó información circunstancial que sugiere
que aún podría existir una pequeña población de león en el MNP y sus zonas circundantes.
Palabras clave: León, , Ghana, Situación, Parque Nacional de Mole
Corresponding author: F. M. Angelici. E–mail: francescomariaangelici@gmail.com
The lion in Ghana:
its historical and current status
F. M. Angelici, A. Mahama & L. Rossi
152 Angelici et al.
Introduction
The rst scientic studies of African lion (
) populations date back to the 1950s and primarily
concern East Africa. Similar studies in West Africa
are almost absent. Generally speaking, it is difcult
to count individual lions (Myers, 1975). They can
cover enormous distances and occupy areas where
they have gone unreported for years, sometimes far
from their normal range of distribution (Smithers,
1983). Much potential information from local people
is likely to remain unknown to science or underesti-
mated, rendering it useless for conservation purposes
(Black et al., 2013). As a consequence, detecting the
presence of a scarce lion population in large areas
is extremely difcult and requires close collaboration
with park staff and local people in order to gather as
much information as possible. According to Henschel
et al (2014), the West African lion
( from Senegal to Nigeria), is currently in serious
danger of extinction (g. 1), with about 400 lions in the
whole of West Africa, probably representing fewer than
250 mature individuals. In the last twenty years, West
African lion populations in Ghana, were estimated at
a few dozen individuals (Wilson, 1993; Chardonnet,
2002; Bauer & Van Der Merwe, 2004) and the species
has recently been considered 'functionally extinct, if
not completely eradicated' (Henschel et al 2010;
Burton et al., 2011a). The lion is 'considered absent
in Ghana' (Henschel et al., 2014) (g. 1). In this pa-
per we aim to reconstruct the historical status of the
lion in Ghana by reviewing the available literature,
unpublished data, and material collected during eld
expeditions in the Mole National Park (MNP) and
other areas between 2005 and 2014.
Materials and methods
Historical data up to 2010
We reviewed the literature on lion distribution and
status in Ghana, including unpublished ofcial reports,
and compared all the available data to identify new
information and any possible inconsistencies in lion
population estimates over the years. We obtained data
from scientic articles and books, park documents,
unpublished reports provided by the Wildlife Division
(Forestry Commission) of the Ghanaian Ministry of Lands
and Natural Resources, IUCN and FAO publications
and rst preliminary expeditions led by the main author
since 2005 to 2009 (Angelici, 2006; Angelici & Petrozzi,
2010). Table 1 lists all the unpublished sources we cited.
Project 'The Pride of Ghana' (Mole National Park,
2011–2014)
The project entitled 'The Pride of Ghana: Local De-
velopment and Assistance Toward for the Sustainable
Management of the Mole National Park and its Fringe
Communities' was ofcially launched in January 2011.
The number of consecutive days spent in the eld to
date was 36 days in 2011, 22 days in 2012, 52 days in
2013 (and 23 days in 2014, in addition to the constant
structured, formal collaboration over the course of the
year with the Wildlife Division staff operating in MNP.
Protocols used during eld activities and habitat
suitability model
From April 12 to August 8, 2011, 20 digital camera
traps (model LTL Welltar 8210A) were positioned.
They were placed in different areas of the park, with
a total of 2,474 trap days and 24 camera stations,
with 1,745 trap days in high–suitability areas (228 on
average) and 501 in low–suitability areas (for a deni-
tion of suitability levels, see below). A regular transect
was not used to position the traps.
In 2012, there were only 163 trap days, using
12 camera traps in 14 different positions. From March
2013 to the present (camera trapping is ongoing), up
to 22 camera traps have been used over the course
of the year, with periods of temporary suspension,
especially during the rainy season, for obvious rea-
sons of accessibility. We have collected thousands of
photographs, which are still under analysis (g. 2 for
all camera positions; in some places, there is more
than one trap position).
The choice of where to position the camera traps
was based on a habitat suitability model (g. 3). This
model was developed on the basis of lion sightings
recorded by park staff between 1968 and 2009 by t-
ting logistic regression models on the habitat features
of lion sightings in MNP. The results of the analysis
(signicant variables) were used to implement the
selected predictive variables in GIS software (ArcGis
v. 9.0) to produce a predictive species suitability map,
to map the areas with the greatest probability
of lion occurrence in the Park. The maps were built
according to three intervals of the probability of occu-
rrence: low–suitability habitat (values from 0 to 0.33);
medium–suitability habitat (from 0.34 to 0.66); and
high–suitability habitat (from 0.67 to 1). The reliability
of the potential distribution model was assessed by
AUC criteria using a Jackknife procedure. The main
positive feature of AUC is its single threshold–inde-
pendent measure for model performance. An AUC
value can be interpreted as the probability that a
presence site, randomly chosen from the dataset,
will have a higher predicted value than an absence
site. The overall model tting was good: AUC=0.886.
Protocol and statistical analyses
We used data recorded over a 41–year time span
by park patrols. Each data entry recorded the con-
rmation of lion presence, including the geographic
coordinates of the site. These data were entered
into DIVA–GIS software. Around each record of lion
occurrence ( = 100), we extracted a circular 100 m
radial buffer, and within this buffer we recorded se-
veral independent variables: (i) linear distance from
the closest road/path (hereby DCR); (ii) rainfall (mm,
per year) (RFL); (iii) linear distance from the closest
pond/water body (DWB); (iv) linear distance from
the closest ungulate prey ( ,
Animal Biodiversity and Conservation 38.2 (2015) 153
A
B
C
D
E
F
N
S
WE
Considered absent
Potentially present
Conrmed present
Ghana border
300 km
, , ,
) (DPR); (v) land use (LNU);
and (vi) elevation (m a.s.l.) (ELE). In addition, we re-
corded the same independent variables for 100 ran-
dom points (also with a 100 m radius) within MNP. A
logistic regression modelling approach was applied
to lion presence/absence (Hosmer & Lemeshow,
1989) using a backward stepwise model (Luiselli,
2006) and the Von Bertalannfy growth function
(Von Bertalannfy, 1934, 1938, 1951, 1964). In these
models, lion presence/absence was the dependent
variable, and the six above–mentioned variables
were the covariates. These techniques are powerful
analytical tools that can analyse the effects of one
or several independent variables, both discrete and
continuous, on a dichotomous dependent variable
(Hosmer & Lemeshow, 1989; Teixeira et al., 2001).
In addition, logistic regression models rely on fewer
statistical assumptions than their alternatives and ge-
nerally produce robust results (Teixeira et al., 2001).
Independence was assessed when 2 < 0.58 (Hosmer
& Lemeshow, 1989; Arntzen & Alexandrino, 2004).
To determine whether the probability of lion pre-
sence in relation to the studied covariates was best
described by backward stepwise logistic regression
or by the Von Bertalannfy growth function, we relied
on a model–selection approach based on the Akaike
Information criterion (AIC) (Burnham & Anderson,
2002) according to the formula:
AIC = −2 log Likelihood + 2K
where n depicts effective sample size, and K is the
number of parameters (= number of variables + 1 to
include the intercept (Sugiura, 1978).
The relative performance of alternative models was
measured using the delta AIC:
∆AIC = AICi – min AIC
where AICi is the AIC value for model i, and min AIC is
the AIC value of the best tting model. Hence, the diffe-
rences between the AIC scores of the various models
(∆AIC) provides a measure of the relative reliability of
the competing models. The advantage of this appro-
ach is that it allows the various competing models to
be ranked according to their relative likelihood and is
not dependent on a threshold value (α–level, Vapnik,
2000). The AIC penalizes the addition of parameters,
and thus selects a model using a minimum number
of parameters according to the principle of simplicity
and parsimony (Akaike, 1973); therefore, the models
with the lowest ∆AIC were selected.
Starting in 2012, we conducted night sessions
(from about 9 pm to 1 am) along some random paths,
listening for any possible lion roars. We conducted two
night sessions in May 2012, 2 in March–April 2013,
and 2 in February 2014.
Fig. 1. Map of West Africa including all protected areas where lion occurrence has been documented
according to Henschel et al. (2014). Protected areas in Ghana: A. Gbele Resource Reserve; B. Mole
National Park; C. Bui National Park; D. Digya National Park; E. Kogyae Strict Nature Reserve; F. Kalapka
Resource Reserve.
154 Angelici et al.
Moreover, as of March 2013, we started to perform
linear transects at night (8.30 pm – 2 am) by car along
the trails in the park using directional headlights at a
constant speed of about 10 km/h. In particular, we
travelled along six transects in 2013, and three in
2014. Each transect was 30–35 km long.
Table 1. A list of the unpublished sources from which they were obtained indirect information, and/or
references were obtained.
Data Area Origin Year
MNP database MNP Wildlife Division (Forestry Commission) 1968–2004
Ghana, compiled by MNP staff
Report by P. J. Pegg MNP FAO library, Rome 1969
on wildlife management
Reports by R. Jamieson Ghana and MNP Wildlife Division 1970–72
(Forestry Commission), Ghana
Reports by Aberdeen MNP Wildlife Division 1974–1977
University Ghana (Forestry Commission), Ghana
expeditions to MNP
Wilson (1993) MNP IUCN library, Gland 1993
Results
Historical data up to 2010
Our ndings from a review of the historical literature
were scattered, sporadic and often inaccurate, testifying
Fig. 2. Localization of camera–trapping sessions in MNP carried out from 2011 to 2014.
N
S
W E
Camera traps 2011
Camera traps 2012
Camera traps 2013
Camera traps 2014
Animal Biodiversity and Conservation 38.2 (2015) 155
to the lack of study of this species in Ghana. This is
possibly because lions were never particularly abun-
dant or widespread in the region. Grubb et al (1998)
meticulously collected several old records of lions from
the late 19th century but even in the map presented
in their paper, it is clear that the last strongholds of
the species were (supposedly) protected areas,
Bui National Park, Digya National Park, GRR, and
MNP, apart from other occasional reports scattered
throughout the North and East of the country on the
Togo border. Lions were even reportedly seen in Ko-
gyae Strict Nature Reserve and Kalapka Resource
Reserve (g. 1), albeit sporadically, at least until the
1990s (Grubb et al., 1998). For an accurate referen-
ce selection, see Grubb et al (1998). Unfortunately,
there are no data on the size of the lion populations
in any area, nor are there any even preliminary data
on their ecology, with the exception of MNP, as we will
see later. In Cansdale’s (1948) provisional checklist
of the Gold Coast, he only mentions the lion as being
present in the open country in the areas of Togoland,
Afram Plains, north–west Ashanti and the Northern
Territories. However, as it is a checklist, he does not
insert any other data, particularly regarding species
abundance or frequency.
Only two reports were found that relate to this
period, and both were very general and based on
rough estimates, not on specic work carried out in
the eld. Mention should be made of a male lion from
Tamale, Doka woodland (about 80 km from MNP),
whose skin is stored in the Natural History Museum
in London, no. 394, dated June 1943 (Rosevear,
1974). To date, Wilson's report (1993), based on
three months of eldwork, is the only document which
includes data on the distribution and ecology of the
species. Wilson conrmed that the lion roar was
heard several times during the survey, in particular
near the headquarters of the Wildlife Division and
along the Lovi River, near Lovi camp (g. 4). Wilson
(1993) claims to have conrmed lion presence in
MNP in at least three different locations. In January
1993, a photo was also taken of a lioness in lacta-
tion (g. 5) by John Grainger, near Gbanwele camp
(g. 4). Various lion droppings were also collected
in at least ve different locations over the 3–month
study, specically in Lovi, Brugbani, Gbanwele,
Samole, and Nyanga (g. 4) (Wilson, 1993). Ran-
gers also collected the skulls of some probable lion
kills. Wilson (1993) also states that 'while the lion
population in MNP is certainly not high there must
be at least sufcient numbers to maintain a breeding
population'. Furthermore, the same author reported
that in December 1992 the rangers saw prides of
up to eight lions all together, including three cubs,
in particular at Lovi and Nyanga. He also concluded
that lions were more easily encountered at Lovi and
Nyanga, along the Lovi River, extending east and
south to Brugbani and Samole. While lions could also
be found going towards Gbanwele and Konkori, as
well as near Kwomwoghlugu. Moreover, they often
Fig. 3. Lion habitat suitability model ( ) in MNP and surrounding areas (for methodology,
see text).
Panthera leo
N
S
WE
Park border
High suitability
Medium suitability
Low suitability
0 20 40 80 120 160 km
156 Angelici et al.
Lion sighting/footprints 2009
Lion sighting 2013
Lion sighting 2014
Roars heard 2011
Roars heard 2012
Roars heard 2013
Roars heard 2014
Buffalo carcassa 2013
Camp
Park H.Q.
Village
Belebile
Holomuni
Ducie
Chasla
Chasia
Camp Sogsima
Camp Ducie
Camp
Gbanwele
Camp
Konkori
Camp
Nyanga
Camp
Dagbori
Camp
Jang
Camp
Nikori
Camp
Jandra
Camp
Grupe
Camp Kuboma
Camp
Kabampe
Camp
Kamanto
Camp Degbere
Camp Semole
Camp
Brugbani
Camp Mognori
Camp
Muruga
Camp
Kwomwoghlugu
Camp
Yazori
Camp
Bawena
Camp
Jang
Soma
Jelinkan
Grupe
Sayire Kobampe
Kananto Palma
Camp
Larabanga
Kong
Daborin
Lovi
Camp
Mognori
Lovi River
Mole River
Belepong
Esalakawu
Camp
Gbantala
Camp
Sabile
Camp
Kulpawn
Camp
Damongo
Muruga
Kaden
Yazori
Bawena
Seriseeni
Camp
Jinfrono
Grubagu
Camp
Grubagu
Wawato
Kporio
Sagiya
Zanwera
Kpulumbo
Park H.Q.
N
S
WE
0 5 10 km
ventured outside the boundaries of the park, accor-
ding to rangers' records, particularly those from the
camps of Kananto, Jang, Gbanwele and Gbantala
(for all localities see gure 4).
In Chardonnet's (2002) account of African lions, he
estimates that 15 (12–18) lions are present in MNP in
the table, while in the text he refers to a range from
15 to 50 lions in MNP according to the estimates of
various specialists. Bauer (2003) and Bauer & Van
Der Merwe (2004) report on an indirect estimate made
by the Ghana Wildlife Society of 20 (12–28) lions in
MNP and 10 (6–14) in GRR.
In 2002, a lioness was killed by poachers in MNP,
and in 2004 a male was shot (g. 6) very close to
the village of Larabanga (g. 4). A few days earlier,
the same lion had killed several heads of cattle and
had also had an aggressive encounter with another
man (see Angelici & Petrozzi, 2010). We collected the
results of questionnaires administered in both MNP
( = 47) and GRR ( = 6), and only 21.3% of the
respondents in MNP reported seeing lions between
2000 and 2009. For further details regarding the
methodology applied, see Angelici & Petrozzi (2010).
According to MNP records for the 1968–2009
period, the maximum number of sightings of a single
lion (of any age) observed on a single occasion was
21, in 1969. No sightings were reported between
1998 and 2008. In 2009, two individuals were spotted
Fig. 4. Detailed map of MNP showing data on lions collected from 2005 to 2014.
Animal Biodiversity and Conservation 38.2 (2015) 157
Fig. 5. Lioness photographed in January 1993 in MNP.
Fig. 6. Lion shot near Larabanga in August 2004.
(Angelici & Petrozzi, 2010). Our results date back to
the end of 2005 when our eldwork was undertaken,
and are intermittent up to 2010. The project was
ofcially launched in 2011. At the same time, the
results of research carried out in MNP by C. Burton
and collaborators (2006–2008), , Henschel et al.
(2010) and Burton et al. (2011a, 2011b), begin to
emerge. During their research, according to their
protocols, the authors did not obtain any ndings
regarding the lion, whereas they collected a lot of
data relating to other mammals. Their conclusion was
that in MNP the lion had likely been functionally, if
158 Angelici et al.
Table 2. Summary of the results obtained in the literature and in the present work (updated to December
2014) regarding lions in Ghana (for all localities see gures 1, 4).
Data Area Reference Year
Several anecdotal records Bui National Park, Digya Grubb et al. (1998) 1893–1960,
of single lion sightings National Park, GRR, MNP, some undated
(late 19th until the 1960s) other occasional reports
scattered throughout the
North and East Kogyae
Strict Nature Reserve and
Kalapka Resource Reserve
General data of occurrence Togoland, Afram Plains, Cansdale (1948) Until 1948
north–west Ashanti and the
Northern Territories
Skin stored in the Natural Tamale, Doka woodland Rosevear (1974) June 1943
History Museum, London, (about 80 Km from MNP)
no. 394 (male)
Many records (sightings, MNP: Lovi, Brugbani, Wilson (1993) 1992–1993
roars, droppings, Gbanwele, Samole, Nyanga,
prey remains, etc.) Konkori, Kwomwoghlugu.
Frequently outside the
park, Kananto, Jang,
Gbanwele, Gbantala (g. 5)
15 (12–18) or 15–50 lions MNP Chardonnet (2002) 2002
(estimate)
20 (12–28) lions in MNP, MNP and GRR Bauer & Van 2004
and 10 (6–14) in GRR Der Merwe (2004)
Lioness shot by poachers MNP, unknown locality Angelici & Petrozzi (2010) 2002
Lion shot by MNP, Larabanga Angelici & Petrozzi (2010) 2004
Larabanga shepherds surroundings (g. 6)
Several direct observations MNP Angelici & Petrozzi (2010) 1968–2009
made by MNP staff
between 1968–2009
(see Discussion)
Questionnaires administered MNP, GRR Angelici & Petrozzi (2010) 2000–2009
in both MNP (n = 47) and
GRR (n = 6), 21.3% in MNP
reported seeing lions, none
in GRR
Sighting of one lion Digya National Park Henschel et al. (2010) October 2008
Case of human–lion Kalapka Resource Reserve Henschel et al. (2010) February 2009
interaction
Sighting of a couple of lions, MNP, Lovi (g. 7) Angelici & Petrozzi (2010) May 2009
footprints, prey remains,
by a park manager
Animal Biodiversity and Conservation 38.2 (2015) 159
not fully, extirpated. The same conclusion was rea-
ched regarding lion presence in GRR. Henschel et
al. (2010), however, did report some anecdotal local
sightings that they considered plausible, although
further conrmation and investigation would be nee-
ded. One such incidence occurred in Digya National
Park where a lone lion was sighted in October 2008
after years with no sightings. An extremely unusual
case regarding a human–lion interaction was reported
in February 2009 in Kalapka Resource Reserve in
south–eastern Ghana near Togo. If conrmed, this
would support what was stated in the introduction of
this article, that a large predator may unexpectedly
'reappear' in areas where it has been declared extinct
despite incomplete knowledge of its status, causing
unforeseen problems (Smithers, 1983; Black et al.,
2013), as recently happened in Gabon (Anonymous,
2015). In May 2009, park staff in MNP, including an
executive manager (Oliver K. Chelewura), clearly
sighted a two lions, a male and a female. This event
was reported in a paper the following year (Angelici
& Petrozzi, 2010). Lion footprints were also observed
(g. 7) at the same sight along with the skull of a
hartebeest that the lions abandoned when they saw
humans. Unfortunately, the picture contains no ele-
ments to estimate footprint size but all the footprints
were found in the general area of the sighting and
appear convincing.
Status 2011–2014
Camera trap sessions
We have not collected any photos of lions to date,
but more than 20 species of mammals have been
photographed, as evidenced in more than 6,000
selected pictures and about one hundred short lms.
Fig. 7. A lion footprint found immediately after
the sighting of two lions in May 2009 in MNP.
Roars heard MNP: staff lodge compound Original data April 2011
Roars heard MNP: along the road Original data May 2012
Mognori–Lovi
Roars heard MNP: three localities (g. 4) Original data April 2013
Sighting of a male lion MNP: close to the Original data August 2013
by a staff guard staff quarters
Possible lion predation MNP: near Brugbani Original data October 2013
of an adult buffalo camp (g. 8)
Roars heard MNP: between Lovi and Original data February 2014
Kwomwoghlugu
A couple of lions sighted MNP: near Gbantala camp Original data August 2014
by poachers, roars heard
Table 2. (Cont.)
Data Area Reference Year
160 Angelici et al.
Reports of direct lion sightings
At 8:07 pm on the evening of April 13 2011, we dis-
tinctly heard a roar from the staff lodge compound.
During the night transect sessions in May 2012, a roar
was heard twice on one occasion and in April 2013,
repeated roars were heard three times in two different
areas within the park (for all roars heard, g. 4). All
these roars were heard at night by one of the authors
(F. M. A.) along with some Ph. D. and MSc. students
and guards Eric Bani and David W. Kabuiri, two of
the most experienced members of the staff.
We have yet to encounter any lions on any night
transects: However, we did encounter and recognized
16 mammal species.
An event of greater note was the sighting by a
guard (D. W. Kabuiri) of an adult male lion very close
to the staff quarters and to the entrance gate to the
park in August 2013. (see g. 4). The following year,
in February 2014, we heard the night roarings of male
lions in the south–central region of the park on two
occasions (g. 4).
In October 2013, we collected some data regarding
the possible lion predation of an adult buffalo. The
buffalo was shot by poachers and retrieved by rangers
and was severely wounded and limping (see the deep
wounds and large, long scratches on the body as well
as the holes on its buttocks made by claws that are
severely infected and full of blowy maggots, g. 8).
In August 2014, we obtained information from a
poacher who was questioned by rangers. The poacher
had seen two adult lions near Gbantala camp (g. 4),
where several roars had also been heard. Further
investigation is currently underway in the region and
camera traps are being placed throughout the area.
All data are summarized in table 2
Discussion
Although compelling evidence has yet to be gathered
( clear videos or photographs), the presence of a
few individual lions in MNP should not be ruled out
. Although the most recent empirical evidence
( a male that was shot) dates back to 2004, the
adult lions observed in 2002 and 2004 could have
reproduced before being killed. Several previous state-
Fig. 8. Buffalo carcass shot by poachers in MNP (October 2013) with clear signs of a probable attacks
by lions, in particular the holes in the hindquarters which are infected and invested with larvae, and the
deep and recent long lateral scratches.
A
B C
Animal Biodiversity and Conservation 38.2 (2015) 161
ments have proven unreliable regarding the extinction
of large cats (Black et al., 2013) and the evidence we
have collected also challenges such a conclusion.
The buffalo shot in October 2013, in addition to the
healed injuries probably caused by other buffalos,
seems to bear the typical signs of a lion attack. Most
of the remaining evidence that has turned up during
our work has been from eyewitness. However, on two
occasions (May 2009 and August 2013) this evidence
was based on reports by qualied and reliable MNP
staff. When management resources are scarce, reports
of a supposedly extinct species can cause controversy
(Roberts et al., 2009), as was the case for lion sightings
in the MNP. Monitoring program of the park has been
criticized by Burton (2012) as not always reliable.
Although often overlooked, the role of paratax-
onomists (local assistants trained by professional
biologists, see Janzen, 1991, 2004) can play a critical
role in conservation (Basset et al., 2004), and infor-
mation provided by trained assistants and the local
people can be as accurate as those of eld biologists
(Danielsen et al., 2014).
The May 2009 sighting is supported by additional
evidence: footprints that can be attributed to a large
cat on the site and the remains of a hartebeest, ap-
parently killed by a large predator. Considering the
eyewitnesses’ statements, the footprints and the typol-
ogy of the prey, the lion is the most likely candidate.
Evidence of lion sightings and roars dated August 2014
near Bantala camp is of particular importance as it
was provided by poachers, who generally understate
the occurrence of wildlife, in particular by not supply-
ing information about lions, for fear of retaliation. The
opinion of Henschel et al. (2010) and Burton et al.
(2011a), which in our view is perhaps too hasty, was
later accepted by many authors and authorities and
reiterated in other articles ( Burton et al., 2011b;
Henschel et al., 2014). Nevertheless, we believe it is
only right to continue to seek out objective data that
attests to the continued persistence of a few lions
in MNP and the immediately surrounding areas. As
pointed out by other authors ( Black & Copsey,
2014), we believe that from the point of view of the
governments and park leaders, a mix of incomplete
knowledge about the presence of a species is better
than assuming its extinction. The possible implications
of a wrongful assumption of the extinction of the lion
would be so important that in the light of indirect evi-
dence we have collected, we believe it is wiser to apply
the Precautionary Principle (Foster et al., 2000) and
assume the survival of the species until more scientic
data tell us the full picture. If there is a chance that
some lions are still present in the MNP, it is essential to
avoid Romeo’s Error (Collar, 1998) for which a species
is thought to be extinct in an area without assessing
all available information. Finally, it should be kept in
mind that considering the lion as extinct quickly leads
to safety rules concerning a direct encounter with the
big cat and all that this implies: danger to livestock but
also danger to man himself, being overlooked. Apart
from the lion attack on a farmer in 2004 (see above),
a good example can be the totally unexpected and
reliable report of a lion in Kalapka Resource Reserve,
Eastern Ghana, towards the border with Togo, where
no–one expected such an occurrence.
In agreement with the proposal by Chardonnet
(2002), we consider the survival of lions in MNP and
in Ghana in general could be of strategic importance to
having a core group of lions forming a central corridor
bridge between Western (Senegal, Guinea) and Eas-
tern (Burkina Faso, Niger, Benin, Nigeria) populations
(g. 1), ideally in the Ivory Coast and in Ghana.
Acknowledgements
We are grateful to Nana Ko Adu–Nsiah, Umaru Farouk
Dubiure, Enoch Amasa Ashie, Oliver K. Chelewura,
(Forestry Commission, Wildlife Division) and all the
MNP staff, who assisted us every step of the way, in
particular Eric Bani and David W. Kabuiri for their eld
assistance. We would also like to thank Gianna Da Re
(Ricerca & Cooperazione NGO, Italy) for her coopera-
tion and assistance during our visits to Ghana. The
institutions sponsoring the project are: the Italian Foreign
Affairs Ministry (DGCS), Rome, Italy; the Department of
Ecological and Biological Sciences, and the Department
of Science and Technology for Agriculture, Forests,
Nature and Energy (University of Tuscia, Viterbo, Italy);
the Forestry Commission (Wildlife Division), Accra,
Ghana; Ricerca & Cooperazione NGO, Rome, Italy. We
also thank Mauro Cella for revising and improving the
English text. We want to thank John Grainger for giving
us a copy of his photo and for allowing us to publish
it in our article. We would also like to thank Damiano
Luchetti, Marco Signore, Dario M. Soldan, Alberto Zilli
and the libraries of IUCN (Gland) and FAO (Rome)
for providing us with documents and publications that
were difcult to nd. Louise Tomsett (Natural History
Museum, London) has provided data on museum
collections. The following people have collaborated
in the eld research: Andrea Caboni, Massimiliano Di
Vittorio, Stefania Gentili, Edoardo Mastrandrea, Fabio
Petrozzi. Luca Luiselli and Massimiliano Di Vittorio
contributed to tting the habitat suitability model. Last,
but not least, many thanks to Emmanuel Do Linh San
for having encouraged us to submit the manuscript, and
for his useful advice, and thanks to the Editor and two
anonymous referees for their useful comments on the
previous draft of this manuscript.
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