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AJE 12829 Dispatch: 10 -11-2020 CE: Karthiyayini B
Journal Name Manuscript No. No. of pages: 6 PE: Raja J.
Afr J Ecol. 2020;00:1–6. wileyonlinelibrary.com/journal/aje
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1© 2020 John Wiley & Sons Ltd
Received: 16 September 2020
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Accepted: 24 October 2020
DOI: 10.1111/aje.12829
SHORT COMMUNICATION
Ant and termite prey of the giant pangolin Smutsia gigantea
Illiger, 1815 in forest–savannah mosaics of Cameroon
Ghislain F. Difouo1,2 | Franklin Simo1,2 | Sévilor Kekeunou1 |
Giovanni Titti Ebangue1 | Lyiong Giscard Ndoh3 | David Olson4
1Labor atory of Zoology of Faculty of Science, Univer sity of Yaound é 1, Yaoundé, C ameroun
2Pangolin Conservation Network, c/o Central Africa Bushmeat Action Group, Yaoundé, Cameroon
3Ministry of Fores try and W ildlife, Bertoua, Camerou n
4WWF-Hong Kong, Man hattan Centre, China
Correspondence: Ghislain F. Difouo, Laboratory of Zoology of Faculty of Science, University of Yaoundé 1, PO Box: 812 Yaoundé, Cameroun.
Email: ghislainfopa49@gmail.com
Funding information
This work w as suppor ted by the Aspire Grants, IDE AWILD, Moham ed bin Zayed Species Conservat ion Fund; Pangolin Conso rtium Grant; and the Rufford
Foundation Grants Program.
1 | INTRODUCTION
Pangolins (Pholidoda: Manidae) primarily feed on ants and termites
(Ashokkumar et al., 2017; Lee et al., 2017; Pietersen et al., 2016). They
consume large amounts of ants and termites with up to 200,000 ants
observed being eaten during a single meal (Shi & Wang, 1985 cited
by Durojaye & Sodeinde, 2014). The diet of Temminck's ground pan-
golin (Smutsia temminckii Smuts 1832) is the most studied of the four
African pangolin species (Coulson, 1989; Kingdon et al., 2013; Pietersen
et al., 2016; Swart et al., 1999). Data on pangolin diets are typically de-
rived from scat (Mahmood et al., 2013) and stomach content analyses
(Ashokkumar et al., 2017; Coulson, 1989; Gao, 1934; Lee et al., 2017;
Minami, 1941) or direct observation of pangolin feeding activities
(Pietersen et al., 2016). Temminck's pangolin demonstrates prey selec-
tivity with preferred ant species in both mesic savannah and arid habitats
(Coulson, 1989; Jacobsen et al., 1991; Pietersen et al., 2016; Swart, 1992).
Diet composition and feeding ecology studies for West and Central
African pangolin species—giant pangolin (S. [Manis] gigantea), white-bel-
lied pangolin (Phataginus tricuspis (Rafinesque 1821)) and black-bellied
pangolin (Phataginus tetradactyla (Linnaeus 1766))—remain limited.
The giant pangolin is the largest and heaviest pangolin species. The
species is listed as Endangered on the IUCN Red List (IUCN, 2019).
It is largely solitary and nocturnal (Bräutigam et al., 1994). The giant
pangolin occurs in primary and secondary rainforest forest, sa-
vannah and forest–savannah mosaic habitats of Central and West
Africa where soils are suitable to search ground dwelling preys
(Nixon et al., 2019). It has been reported to feed predominantly on
five termite genera, including Macrotermes, Cubitermes, Apicotermes,
Protermes and Pseudacanthotermes, and two ant genera, Palthothyreus
and Anomma (Kingdon, 1972; Nixon et al., 2019). Eleven ant species
was also recorded previously as part of the giant pangolins diet (see
Bequaert, 1922, cited by Hoffmann et al., 2019). The specific prey spe-
cies and their relative proportion consumed are not described. Here,
we identify and quantify ant and termite species found in stomach
content and scat of giant pangolins from an ecotone of Cameroon and
compare to the community and abundance profile of cursorial ants of
this area to evaluate prey selectivity.
2 | STUDY AREA
The survey was conducted in Mpem et Djim National Park (MDNP)
(Figure 1). Mpem et Djim National Park (5°–5°20' N/ 11°30'–12° E;
976 km2; average altitude of 640 m) is located in the Central Region
of Cameroon in the Mbam and Kim Division. The protected area is lo-
cated in a forest–savannah transition (ecotone) with a mosaic of forest
habitats and large block of forests (Dames & Moore, 1999). This park
is characterised by classic Guinean climate with four seasons annu-
ally; the mean annual rainfall ranges between 1,800 and 2,000 mm
per year, while the annual temperature averages 22–29°C (Tsalefac
et al., 2003). Both park vegetation formations are forest and savannah
(Figure 1) with several habitat types including mosaic of closed-canopy
and open-canopy secondary forests, woodland and grassland savan-
nahs, and gallery forests.
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3 | SURVEY METHODS
3.1 | Giant pangolin stomach content and scat
analysis
During a community survey in the villages surrounding Mpem et
Djim National Park (Difouo et al., 2020), we found a dead juvenile
giant pangolin. The Ministry of Forestry and Wildlife (MINFOF) pro-
vided a research permit N°0805 that allowed stomach contents of
the pangolin to be examined. The stomach content was collected
and stored in 90% ethanol and, just prior to analysis, was soaked in
fresh water and dried for 10 min on blotting paper following protocol
from Mahmood et al. (2013).
One fresh scat sample (estimated as < 24 hr old) was collected in
the Mpem et Djim grassland savannah. We assume the scat belonged
to an adult giant pangolin because: (a) its large dry weight (120.1 g)
precludes all small-size myrmecophagous, other pangolin species
and juvenile giant pangolin and the scat appeared different in shape
(fusiform) and length (>8 cm) from that described for an aardvark
(Orycteropus afer Pallas 1766) scat (Chame, 2003); (b) our local guides
expressed confidence the scat was from giant pangolin and not an
aardvark; (c) camera-trap images from the burrow captured only giant
pangolins utilising the burrow and no aardvark (Simo et al. in prep); and
(d) the relatively close similarity of the prey species composition be-
tween the purported scat and the giant pangolin stomach content (see
Results). The scat sample (n = 1) was collected and stored in a plastic
Ziploc bag prior to analysis following previously published methods
(Mahmood et al., 2013).
3.2 | Species and genera identification
Following Lee et al. (2017), insects found were sorted into mor-
phospecies based on ex ternal morphological characteristics of
FIGURE 1 Locations of the surveyed sites in the MDNP in Central and East Regions, Cameroon (source: National Institute of
Cartography INC 2012 modified)
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COLOUR
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DIFOUO e t al.
entire individuals and fragments, including heads and thorax that
are strongly keratinised. Ant morphospecies groups were identified
using the dichotomous keys in Hölldobler and Wilson (1990), Bolton
(1994), and the African Ants systematic database (www.antba
se.org). The morphometry of the head and position of the eye on
the head among the ant species were used to identify them to spe-
cies level based on Bolton (1994). Termites were identified based on
soldier castes, identified to genus and species, where possible, using
appropriate dichotomous keys (Bouillon & Mathot, 1965; Emerson
et al., 1928; Ruelle, 1970; Sands, 1965). Specific richness and rela-
tive abundance of ants and termites were recorded. Only heads
were used as counting unit to prevent errors arising from repeated
counting of body fragments. The ant prey species selectivity of the
scat and stomach content samples was evaluated by the equation of
Manly and colleagues (1993).
4 | RESULTS
4.1 | Ant prey
The juvenile giant pangolin stomach content yielded 1,027 ants (71%
of insect s recorded; Table 1) comprising nine species. Large ant spe-
cies predominated, mostly Camponotus brutus Forel 1886 (515 indi-
viduals , 35%; Table 1) and Palthothyreus tarsatus (Fab ric ius 1798) (24 0
individuals, 17%). The faecal sample had 3,127 ants (88% of insects
recorded), belonging to eight ant species. Large and medium-sized
ant species were common, such as Polyrhachis militaris Fabricius
1781 (2,171 individuals, 61%; Table 1) and the larger C. brutus (538
individuals, 15%). Differences in the composition of the faecal and
stomach content samples may simply reflect each pangolin's latest
feeding event, rather than individual or age-related preferences. In
both, relatively large ant species were most common in the two pan-
golin's diet as sampled here, namely C. brutus and P. tarsatus.
The giant pangolin stomach content and scat samples suggest
that both pangolin individuals had fed on similar species (six com-
mon species representing more than 55% of their diets). Despite
their similar composition of species (see Tables 1 and 2), the stom-
ach content did not closely match that found in the scat sample in
terms of relative abundance. Polyrhachis militaris represented 61%
of ants recorded in the pangolin scat compared to 0.5% in the pan-
golin stomach contents, where C. brutus dominated (35%). This may
simply reflect the ant species each individual most recently fed on.
We compared the giant pangolin stomach and scat content ant com-
munities with the ant communities from each habitat of MDNP. The
ant P. militaris was primarily collected in grassland savannah (GS)
and woodland savannah ( WS), while C. brutus was most abundant in
near-primary forest (NPF) and secondary forest (SF) (Table 1).
4.2 | Termite prey
Termite prey were less than 30% of the total inver tebrates recorded
in both the scat and the stomach content . The stomach content
TABLE 1 Absolute and relative abundance (RA) and prey selectivity index (Ẇ) for ants between the stomach (n = 1) content and scat
(n = 1) of two different individuals of giant pangolin
Ant species
GP-Scat (Adult) GP-Stomach (Juvenile)
Body Avg.
length (mm) RA Habitat# Ind (%)
Selectivity
index (Ẇ) # Ind (%)
Selectivity
index (Ẇ)
Camponotus brutus 538 (15) 34.3*** 515 (35) 80.66*** 15.64 ± 0.64 0.44 NPF
Camponotus flavomaginatus 120 (3) 1.41*147 (10) 4.24** 9.2 3 ± 0. 52 2.39 SF
Camponotus maculatus - - 22 (2) 0.54*12. 0 ± 0.35 2.79 SF
Cataulacus wessi 150 (4) 421*** - - 7. 27 ± 1.72 0.01 NPF
Odontomachus troglodytes 66 (2) 0.58 2 (<1) 0.04 10. 50 ± 1.92 3.17 NPF
Palthothyreus tarsatus 7 (<1) 0.07 240 (17) 5.93** 15.55 ± 0.78 2.79 NPF
Polyrhachis laboriosa (F. Sm ith) - - 27 (2) - 5.38 ± 1.47 - -
Polyrhachis militaris 2 ,171 (61) 82.27*** 8 (1) 0.74*6. 27 ± 1.72 0 .74 GS, WS
Tetramorium aculeatum 4 (<1) 0.01 - - 2.17 ± 0.23 8.34 NPF
Pheidole sp.1 - - 2 (<1) 0.03 2.38 ± 0.44 5.57 NPF
Pheidole sp.2 71 (2) 0.29 63 (4) 0.63*2.12 ± 0.22 6.94 NPF
Tot al 3,127 (88) 1,026 (71)
Note: Average length (mean ± standard error (SE)) of ant body and relative abundance of each ant species per habitat type sampled in the MDNP (see
Difouo et al., in prep). The grey colour in rows denotes the impor tance of the species in the diet composition of pangolin; Habitat type codes are GF,
Gallery forest, NPF, near-primary forest, WS, Woodland savannah, Sl, Saltworks, GS, Grassland savannah, Sw, Swamp, SF, Secondary forest. Dashed
cells denote where information was not provided.
*Lower selectivity.
**Higher prey selectivity.
***Denotes a highest prey selectivity.
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yielded 423 termites (29% of insects recorded) comprising six ter-
mite species (Table 2). The scat had 439 termites (12% of insects re-
corded) comprising four species. Pseudacanthotermes militaris Hagen
1858 was the most abundant termite species in both pangolins diet
composition (313 [22%] of invertebrates recorded in stomach con-
tent and 312 [9%] of inver tebrates recorded in scat) followed by
Macrotermes bellicosus (Smeathman 1781) (89 individuals [6%] in the
stomach contents and 74 individuals [2%] in the scat; Table 2). Both
termite species are the largest species sampled in the scat, stomach
contents and from habit ats in the protected area (see Difouo et al.,
in prep). The most abundant termite species fed upon by both giant
pangolins, P. militaris, was sampled most in savannah habitat s.
5 | DISCUSSION
Although based on initial and limited data, our survey confirms that
giant pangolins eat ants and suggest s they prefer relatively large
ants (>15 mm length) and, in general, feed less on small species de-
spite their greater abundance. The two giant pangolins sampled fed
on eight to nine ant species, respectively, with eleven ant species in
total in their collective samples similar to Bequaert (1922). Pietersen
and colleagues (2016) recorded five species in Temminck's pangolin
diet and Swart and colleagues (1999) recorded fifteen. Camponotus
brutus and Palthothyreus. tarsatus were the most abundant ant spe-
cies in the juvenile giant pangolin's stomach, while Polyrhachis.
militaris and C. brutus predominated in the adult pangolin's scat.
Again, this difference may simply reflect their latest feeding events.
Palthothyreus recorded in this survey were mentioned by Kingdon
et al. (2013), and Tetramorium, Camponotus, Polyrhachis and Pheidole
by Bequaert (1922) as part of the giant pangolins diet, while other
genera recorded here are not previously recorded. None of ant
species recorded was common to the giant pangolins diet examined
in previous studies except Tetramorium aculeatum (Mayr…). Variation
in individual preferences, and spatial and seasonal availability of prey
may account, in part, for differences in prey species obser ved among
studies (Swart et al., 1999).
The primary ant prey species of the giant pangolins, namely
Camponotus brutus (within the stomach of the juvenile pangolin) and
Polyrhachis militaris (within the scat of the adult pangolin), consti-
tuted < 1% of the overall species composition (i.e. number of spe-
cies recorded in cursorial ant surveys) of the surveyed area (Difouo
et al., in prep), although making up 35% and 60% of ants consumed,
respectively. This suggests that the giant pangolins preferentially
feed on cer tain prey rather than feeding on the most abundant
ant species. Similar behaviour is reported for Temmincks’ pangolin
(Pietersen et al., 2016; Swart et al., 1999) which favours Anoplolepis
custodiens (Smith 1858). Larger ants appear to be preferred by giant
pangolin, which may increase their foraging efficiency, in terms of
time, energy and nutrient value (Swart et al., 1999). The nutritional
value of ants increases with larger body size, greater population den-
sity and larger nest structure (Swart et al., 1999). We examined an
adult pangolin scat and stomach content from a juvenile pangolin;
it remains uncertain if juvenile giant pangolins forage in same ways
as the adults as has been observed in the Temmincks’ pangolin diet
(Pietersen et al., 2016).
The surveyed giant pangolins consumed larger termite spe-
cies (>10 mm in length) and, in general, do not feed much on
smaller termites despite their greater occurrence. Among the 53
termite species recorded in MDNP (Difouo et al., in prep), four
and six termite species were identified in the scat and stomach
contents, respectively. The termite species found in both stom-
ach and scat samples are different to those found in previous
studies, which focused on different pangolin species (Pietersen
et al., 2016; Swar t et al., 1999). Among the prey species recorded
and being eaten by giant pangolin here, the genera Macrotermes,
Pseudacanthotermes and Cubitermes were previously mentioned by
Kingdon and colleagues (2013) and Nixon and colleagues (2019).
Pseudacanthotermes militaris, the most abundant termite species
in both scat and stomach samples, has similarly larger body size as
the termite species recorded by Swart and colleagues (1999) and
are larger than the species recorded by Pietersen and colleagues
(2016) for the Temminck's pangolin. Such as for ants, giant pango-
lin may favour eating larger termite species for increased foraging
efficiency (Swart et al., 1999). Termites represent less than 30%
of the examined giant pangolin diets, while ants were up to 80%
of the insec ts recorded. Pangolins may favour ants over termites
(Coulson, 1989; Pietersen et al., 2016; Swart et al., 1999). It re-
mains uncertain if termites are eaten by giant pangolin for com-
pensatory nutrients or antidiarrhoeal role as suggested for other
mammal species (Deblauwe, 2009).
ACKNOWLEDGEMENTS
We thank the Zoological Society of London–Cameroon for techni-
cal assistance. We appreciate the support of MINRESI (Ministry of
TABLE 2 Relative abundance (R A) for termites between the
stomach (n = 1) content and scat (n = 1) of two different individuals
of giant pangolin
Termite species
GP-Scat
(Adult)
GP-Stomach
(Juvenile)
Body Av.
length (mm)
Number of individuals
(%)
Ancistrotermes crucifer - 6 (<1) 2.01 ± 0.01
Cubitermes sp. -13 (<1) 7.31 ± 0.12
Macrotermes bellicosus 74 (2) 89 (6) 14.3 4 ± 0.42
Macrotermes lilljeborgi 7 (<1) 016.7 ± 0.56
Pseudacanthotermes
militaris
312 (9) 313 (22) 10.05 ± 0.12
Macrotermes sp. 46 (1) 2 (<1) -
Termitidae sp. - 2 (<1) -
Tot al 439 (12) 425 (29)
Note: Average length of termites (mean ± SE). The grey shading
denotes the two most abundant taxa sampled and dashed cells where
information was not provided.
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Scientist Research and Innovation) for the research authorisation
and the MINFOF (Ministry of Forestry and Wildlife) for the research
permit. We thank the Conservators Menvi Abessolo I. Charles from
DDNP and Bisseck J. Pierre from MDNP for their ongoing support
for this research and logistical support. We acknowledge Oma T.
Diane and Dr. Wandji A. Christel for their help with data input and
assisting in data analysis. We thank Dr. Nzoko Fiemapong A. Richard
and Dr. Darren Pietersen for their comment s and edits on earlier
drafts. We are very grateful to the village chiefs and field team for
permitting us to work with them.
CONFLICT OF INTERESTS
The author(s) declared no potential conflicts of interest with respect
to the research, authorship and/or publication of this article.
DATA AVAILA BILIT Y S TATE ME NT
The data that support the findings of this study are available from
the corresponding author upon reasonable request.
ORCID
Ghislain F. Difouo https://orcid.org/0000-0002-7905-6538
Franklin Simo https://orcid.org/0000-0002-2607-9648
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