Chapter
Diet and Ecology of Extant and Fossil Wild Pigs
3 Antoine Souron
Introduction
Most extant wild pig species (Suidae) are omnivorous and con-
sume a large variety of foods, including animal matter, plant
matter (leaves, grass, forbs, ferns, fruits, nuts, roots), fungi,
faeces, and even soil (e.g. Meijaard et al. 2011). Species of the
genera Babyrousa, Sus, and Potamochoerus are omnivorous and
comprise the large majority of the specific diversity containing
13 of 17 species recognized by a recent synthesis (Meijaard et al.
2011).
Extant omnivorous suids all display similar cranio-mandibular
and dental morphologies (see below) adapted to process a wide
array of food items. Most extinct suids display morphologies simi-
lar to those of extant omnivorous suids. By analogy, they are usually
considered to be omnivorous.
I will focus here on the exceptions within the Suidae family
and review the morphological and ecological data that docu-
ment how some species of African suids became adapted to
more specialized diets dominated by plant matter, and espe-
cially grasses. Herbivorous suids belonging to the subfamilies
Tetraconodontinae and Suinae in Africa diversified during the
last 10 million years. The extant giant forest hogs (Hylochoerus)
and warthogs (Phacochoerus) are the only relics of those
radiations, being descendants of the Plio–Pleistocene gen-
era Kolpochoerus and Metridiochoerus, respectively (Harris &
White 1979; Souron et al. 2015a).
Climatic and Environmental Background
Strong climatic changes occurred during the Neogene and
Quaternary periods with a global trend to a cooler global climate
and a drier climate in tropical Africa (e.g. deMenocal 2004). The
expansion of tropical grasslands dominated by grasses using
the C4 photosynthetic pattern is well documented based on
stable carbon isotopes of paleosols and mammal teeth, as well
as paleobotanical remains (Bonnefille 2010; Uno et  al. 2011;
Cerling et al. 2015).
Roughly synchronous with those climatic and vegetation
changes, dramatic changes in the faunas are also documented,
notably in the rich fossil record of eastern Africa. Various mam-
mals with strong dental adaptations to the consumption of abra-
sive grasses (hypsodonty, megadonty, etc.) become more and
more abundant as the landscapes changed. The African suids
form a part of this diversification and they started exploiting C4
grasses during the late Miocene, after most other groups of her-
bivorous mammals (Uno et al. 2011). The details of the faunal
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changes are subject to much debate (continuous replacement in
Bibi & Kiessling 2015 versus pulse turnovers in other studies).
Taxonomic and Phylogenetic Framework
Due to the rampant convergent evolution towards similar
dental morphologies and the fragmentary nature of the first
fossils discovered, the species-level taxonomy of the African
Tetraconodontinae and Suinae is still much debated (e.g.
Boisserie et al. 2014; Souron et al. 2015a).
The genus Nyanzachoerus first occurred in Africa some time
during the middle Miocene (following a dispersal event from a
Eurasian ancestor) and became abundant after 8 Ma with at least
eight species recognized by a recent study (Boisserie et al. 2014).
Around 4 Ma, Nyanzachoerus gave rise to the more derived genus
Notochoerus, represented by at least four species (Figure 3.1).
The genus Kolpochoerus first occurred in Africa around
5.5 Ma and around eight species are known (Souron 2012;
Souron et al. 2015a). It was first represented by a single lineage
(divided into two to three chronospecies) during the Pliocene,
followed by a modest radiation giving rise to six species dur-
ing the Pleistocene (Figure 3.2). Kolpochoerus gave rise to the
extant genus Hylochoerus sometime during the Pleistocene, but
the fossil record of the latter is extremely scarce (Souron 2012;
Souron et al. 2015a).
The genus Metridiochoerus first occurred in Africa by at least
3.4 Ma (White et al. 2006) and possibly earlier, by 3.8 Ma (Souron
2012). There was a modest radiation during the Pleistocene
(around seven species) and it gave rise to the extant genus
Phacochoerus. Again, the fossil record of the latter is scarce and
it is difficult to decipher the timing of the origin of Phacochoerus
(Figure 3.3).
Methodology
Most paleoecological reconstructions for extinct African suids
are based on: (1) ecomorphology of teeth; and (2) stable carbon
isotopes of the enamel.
Ecomorphology
Ecomorphology uses the relationship between the ecology and
morphology of extant mammals as a guide to understand the
ecology of extinct mammals. The evolutionary history of African
suids left us with three extant genera with extreme ecologies and
morphological adaptations, encompassing most morphological
disparity found in their extinct relatives.
29
 Part I: Evolution, Taxonomy, and Domestication

Figure 3.1  Changes in third molar morphology and stable carbon isotopes in Nyanzachoerus and Notochoerus. Right panel: chronological ranges for each species
(grey bars). First and last occurrences follow White (1995), Boisserie et al. (2014), and personal observations. Abbreviations: syr, Ny. syrticus; tul, Ny. tulotos/Ny. devauxi; khi,
Ny. khinzir; dev, Ny. devauxi; way, Ny. waylandi; aus, Ny. australis; kan, Ny. kanamensis; jae, Ny. jaegeri; eui, No. euilus; cap, No. capensis; sco, No. scotti; cla, No. clarki. A selec-
tion of upper third molars in occlusal and lateral views illustrate the morphological changes and the omnivorous versus herbivorous morphotypes. Left panel:
stable carbon isotopes of enamel of Nyanzachoerus and Notochoerus (personal compilation of literature, full list of references available on request).

The extant species of Potamochoerus are typical omnivorous
suids. The premolars and incisors are well developed. The third
molars are anteroposteriorly short, low-crowned, and display
few main cusps with thick enamel and bunodont morphology
(Harris & White 1979). The insertions for the temporal mus-
cle are well marked on the cranium. The insertions of the ros-
tral muscles controlling the movements of the rhinarium when
digging for underground foods are also well marked. Those
characteristics are also found in extant species of the mostly
Eurasian genus Sus.
The two other genera of extant African suids (Hylochoerus,
Phacochoerus) display much more specialized morphologies
related to their herbivorous diets. Their premolars are reduced
30
both in terms of number and size. The incisors are also reduced or
even absent. The third molars are anteroposteriorly more elon-
gated, higher crowned, and display numerous main cusps with
thinner enamel (Souron 2012). The insertions for the temporal
and rostral muscles are reduced, whereas the insertions for the
pterygoid and masseter muscles (involved in the lateral compo-
nent of the mastication) are strongly marked. All those morpho-
logical characteristics are presumably related to the functional
needs to reduce tough plant matter to small fragments.
In Hylochoerus, the molars are characterized by well-separated
and anteroposteriorly compressed cusps, giving a bunolopho-
dont occlusal surface (Souron et al. 2015b). The cusps are encom-
passed within a thick cover of cement. The third molars are

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 Chapter 3: Diet and ecology of extant and fossil wild pigs

Figure 3.2  Changes in third molar morphology and stable carbon isotopes in Kolpochoerus and Hylochoerus. Right panel: chronological ranges for each species
(grey bars). First and last occurrences follow White (1995), Souron (2012), Souron et al. (2015a), and personal observations. Abbreviations: deh, K. deheinzelini; afa,
K. afarensis; lim, K. limnetes; old, K. ‘olduvaiensis’ (in need of revision); pai, K. paiceae; coo, K. cookei; phi, K. phillipi; maj, K. majus; mei, H. meinertzhageni. A selection of
upper third molars in occlusal and lateral views illustrate the morphological changes and the omnivorous versus herbivorous morphotypes. Left panel: stable
carbon isotopes of enamel of Kolpochoerus and Hylochoerus (personal compilation of literature, full list of references available on request).

Figure 3.3  Changes in third molar morphology and stable carbon isotopes in Metridiochoerus and Phacochoerus. Right panel: chronological ranges for each spe-
cies (grey bars). First and last occurrences follow White (1995), Souron (2012), and personal observations. Abbreviations: Met, Metridiochoerus sp.; sha, M. shawi; pha,
‘Sus’ phacochoeroides (needs to be transferred into Metridiochoerus); and, M. andrewsi; com, M. compactus; hop, M. hopwoodi; mod, M. modestus; Pha, Phacochoerus
sp.; afr, Ph. africanus; aet, Ph. aethiopicus. A selection of upper third molars in occlusal and lateral views illustrate the morphological changes and the omnivorous
versus herbivorous morphotypes. Left panel: stable carbon isotopes of enamel of Metridiochoerus and Phacochoerus (personal compilation of literature, full list of
references available on request).
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 Part I: Evolution, Taxonomy, and Domestication
moderately elongated with main cusps that are more numerous
than in Potamochoerus. The third molar crowns are also moder-
ately heightened. Those features are related to the versatile her-
bivorous diet of Hylochoerus, comprising grasses and other kinds
of plants (e.g. d’Huart 1978).
In Phacochoerus, the occlusal surfaces of the molars are flat.
The narrow, elongated, and very high-crowned third molars are
composed of numerous cusps, all tightly packed together within
a cover of cement. Those features are related to the specialized
diet of Phacochoerus, comprising almost only grass (both aerial
and underground parts) (Vercammen & Mason 1993).
The morphologies and ecologies observed in extant African
suids are therefore used as a crude guide to reconstruct the diets
of extinct relatives. The third molars, frequently fossilized, are
especially useful to classify the extinct suids into omnivorous
morphotypes (similar to Potamochoerus) and herbivorous
morphotypes (similar to Hylochoerus and Phacochoerus). This
approach was the most commonly used before the develop-
ment of other proxies such as stable carbon isotopes (Cooke &
Wilkinson 1978; Harris & White 1979; Kullmer 1999).
Stable Carbon Isotopes
Stable isotope biogeochemistry uses relative abundances of dif-
ferent isotopes of the same element preserved in the animal tis-
sues to reconstruct its ecology and environments (e.g. Cerling
et al. 2010). I give an overview of the use of stable carbon iso-
topes in paleoecology and its limitations before presenting the
results of the literature review.
Stable isotopes are non-radioactive atoms of the same chem-
ical element that differ by their number of neutrons. Carbon
has two stable isotopes, the ‘light’ 12C (six protons, six neutrons)
and the ‘heavy’ 13C (six protons, seven neutrons). Due to their
slightly different mass, light and heavy isotopes behave dif-
ferently in their geochemical cycles (isotopic fractionation).
During a change of physical state, one of the isotopes reacts less
than the other, which gives different isotopic ratios in the prod-
ucts. These phenomena of fractionation are used to track differ-
ent elements in the cycles.
The tiny quantities of the different stable isotopes are meas-
ured using an isotope-ratio mass spectrometer (IRMS) and
expressed as parts per thousand (‰) relative to a standard
defined internationally. δ13C is reported as a difference com-
pared to the international standard V-PDB (‘Vienna Pee Dee
Belemnite’) and is calculated as follows:
δ13C = [(RSAMPLE/RSTANDARD − 1)] * 1000
with R = ratio of heavy to light isotope.
Dental enamel, due to its strong mineralization and low
porosity, is less prone to chemical exchanges during diagenesis
(Wang & Cerling 1994), and it is therefore the favored target for
isotopic analyses of fossils. Enamel is not reworked during the
lifetime of one individual and environmental variations (cli-
mate, diet) are recorded during mineralization. Enamel miner-
alization is a complex, discontinuous process that takes place
in several stages and involves several successive mineralization
fronts running in different directions. Thus, no matter the size
of the enamel sample, the resulting isotopic ratio represents
32
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an average based on a time period (Balasse 2002). That can be
important when comparing data coming from different sam-
pling strategies.
δ13C recorded in animal enamel reflects that of the food
items that are mainly consumed during the dental growth. The
isotopic enrichment factor ε expresses the isotopic fractionation
and the difference between the two phases (enamel and food).
It is defined as follows (see Cerling & Harris 1999 and Passey
et al. 2005):
εenamel-food = (((1000 + δ13Cenamel)/(1000 + δ13Cfood)) − 1) * 1000 .
Knowing εenamel-food, it is possible to estimate δ13Cfood from
δ Cenamel. That variable was estimated through experimenta-
13
tions. Most studies indicate a value of ε close to 14 ‰ for large
herbivores (Cerling & Harris 1999; Passey et al. 2005; Warinner
& Tuross 2009). The ε factor varies according to the digestive
physiology of the animal (Passey et al. 2005), the digestibility
of consumed food items (Codron et al. 2011), and even other
factors not well understood that could be linked to stress or the
well-being of the animal.
In suids, the results indicate ε values ranging from 12.3 ‰
and 13.3 ‰ for Passey et al. (2005) depending on the digestibil-
ity of the food items (respectively, pure C3 versus pure C4), and
between 14.4 ‰ and 14.6 ‰ for Warinner and Tuross (2009) for
a similar diet.
For that synthesis, I use as ε the average between the mini-
mal and maximal values from Passey et al. (2005) and Warinner
and Tuross (2009); that is, 13.3 ‰. Isotopic composition of
consumed food items can be estimated from that of enamel as
follows:
δ13Cfood = 1000 + δ13Cenamel * (1000/(εenamel-food + 1000)) .
δ13C of terrestrial plants consumed by mammals depends
mostly on the photosynthesis type used (C3, C4, and CAM, or
Crassulacean Acid Metabolism). Those types of plants do not
fractionate the stable carbon isotopes of the atmospheric CO2 in
the same way and their tissues display different δ13C values. C3
plants display δ13C values ranging from –38 ‰ to –22 ‰ (modal
value –27 ‰), whereas C4 plants display δ13C values ranging
from –19 ‰ to –9 ‰ (modal value –12.5 ‰). CAM plants display
intermediate δ13C values (Farquhar et al. 1989; Kohn 2010).
In modern tropical and subtropical Africa, in low-altitude
areas, all trees as well as most shrubs and bushes (dicotyledons)
are C3 plants, whereas 90 per cent of grasses (monocotyledons)
are C4 plants (Livingstone & Clayton 1980; Sage & Monson
1999). Those two types of plants correspond broadly to the
plants consumed by browsers and grazers. As δ13C values of C3
and C4 plants do not overlap, it is possible to distinguish brows-
ers from grazers in tropical environments from Africa based on
enamel δ13C values. This rationale is used in most studies deal-
ing with the paleoecology of Neogene and Quaternary African
mammals (see syntheses by Cerling et al. 2010, 2015).
At low altitudes in tropical environments, C3 grasses are rare
and limited to humid and shady habitats. At higher altitudes, C3
grasses are abundant. Above 1300 m, their abundance increases
and they are the only grasses present above 4000 m (Livingstone
& Clayton 1980). Similarly, in non-tropical African habitats,

and in higher latitudes in other continents, C4 grasses are very
rare. In that case, the browser/grazer dichotomy is not detect-
able using stable carbon isotope biogeochemistry.
Additional factors also complicate the use of that C3 browser/
C4 grazer dichotomy in paleoecology, even in low-altitude
tropical environments in Africa.
1. CAM plants, with their intermediate δ13C values, are usu-
ally neglected in paleoecology as they form a negligible
part of the vegetal biomass targeted by African herbivores.
They mostly include xerophytic plants (cactus-like),
succulent plants from arid habitats, and epiphytic plants
(liana-like) from closed forested environments (Sage &
Monson 1999). Those plants are, however, consumed by
suids, for example by common warthogs in dry environ-
ments (Vercammen & Mason 1993) or giant forest hogs in
closed environments.
2. Sedges (Cyperaceae) are plants usually inhabiting humid
environments, notably numerous kinds of reeds. They use
varied photosynthesis types. Sedges are consumed abun-
dantly by suids (Meijaard et al. 2011).
3. Aquatic plants display extremely variable δ13C values that
rather depend on physical and chemical characteristics
of the aquatic habitat occupied: pH, quantity of dissolved
organic matter, and speed of flow (Sage & Monson 1999).
All extant suids, and notably Potamochoerus, are known to
consume at least some aquatic plants (Meijaard
et al. 2011).
4. δ13C values of plants also show seasonal variations. C3
plants are strongly influenced by seasonal environmental
variations (water stress, light intensity/brightness) and their
δ13C display seasonal variations, usually around 2 ‰ to 3 ‰
(Smedley et al. 1991). Seasonal variations of C4 plants seem
to be weaker, less than 2 ‰ (Codron et al. 2005).
5. δ13C values also vary within plants among the different
organs. Non-photosynthetic tissues of C3 plants (ligneous
stems, roots, seeds, and fruits) display higher δ13C values
than those of leaves (Cernusak et al. 2009). The difference
between δ13C values of roots and leaves was estimated to
be around 1–3 ‰ for several C3 plants. Similar values are
estimated for the difference between seeds and leaves.
Intraplant variation in C4 plants is weaker, lower than
1 ‰ (Cernusak et al. 2009). Extant omnivorous suids
consume a lot of roots and fruits of C3 plants (Meijaard
et al. 2011).
6. All extant suids consume at least some animal matter
(Meijaard et al. 2011). Animal tissues display very differ-
ent δ13C values depending on the organs consumed (up
to 6 ‰ between two organs according to Gearing 1991).
Consumption of carrion by suids can therefore introduce
one supplementary source of variation of δ13C values of
their enamel.
7. It is also necessary to take into account the relative propor-
tions of C3 and C4 herbaceous plants within the herbaceous
stratum of the Neogene and Quaternary environments.
Although the modern situation is relatively simple, with
C4 grasses strongly dominating the C3 herbaceous plants in
low-altitude tropical environments, the situation was very
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Chapter 3: Diet and ecology of extant and fossil wild pigs
likely much more complicated in the past. The timing and
rhythm of the transition from C3-dominated grasslands to
C4-dominated grasslands are totally unknown. C4 grasses
are clearly abundant since the late Miocene (e.g. Cerling
et al. 2010; Uno et al. 2011). However, it is possible that
C4 grasses were not that dominant as today, with a more
mosaic pattern (C3 and C4 herbaceous plants) during the
Pliocene and the beginning of Pleistocene (Bonnefille
2010). For example, based on studies of phytoliths,
Rossouw and Scott (2011) showed that the herbaceous
stratum at Laetoli during the late Pliocene was composed
of a combination of C3 and C4 herbaceous plants. Similarly,
several studies of paleosols δ13C indicate that relatively low
values corresponding to C3-rich environments (around
–8 ‰) were common up to 2.7 Ma whereas grass pollens
were abundant (Bonnefille 2010). That would be congruent
with a greater relative abundance of C3 plants in the herba-
ceous stratum during the Pliocene.
When interpreting stable carbon isotopic data from fos-
sils, it is common to use the dichotomy C3 browser/C4 grazer
observed in modern low-altitude tropical environments. This
dichotomy was likely much more blurred during the Neogene
and Quaternary. The tendency by paleontologists to interpret
the percentage of C3 plants in the diet of fossil mammals as
representing non-herbaceous plants could therefore lead to an
underestimation of the proportion of herbaceous plants con-
sumed (e.g. Harris & Cerling 2002; Cerling et al. 2015).
By adding the value of isotopic enrichment factor εenamel-food
(13.3 ‰) to the modal values of the δ13C of C3 (–27 ‰) and C4
plants (–12.5 ‰), the threshold values of enamel δ13C corre-
sponding to diets composed of 100 per cent of C3 plants and 100
per cent of C4 plants are obtained. I consider that a suid feeding
exclusively on C3 foods has δ13C values lower than –13.7 ‰ and
that a suid feeding exclusively on C4 foods has δ13C values higher
than 0.8 ‰.
As the atmospheric δ13C has greatly increased since 1850
due to anthropogenic activity (the ‘fossil fuel effect’), it is nec-
essary to apply a correction of 1.4 ‰ when comparing δ13C of
fossil specimens to those of extant specimens (Francey et  al.
1999). The 100 per cent C3 and 100 per cent C4 thresholds of δ13C
enamel of fossil suids are, respectively, –12.3 ‰ and +2.2 ‰. In
every figure that combines extant and extinct specimens, δ13C
values of extant specimens are shifted by 1.4 ‰ so that they can
be directly compared to those of fossil specimens.
Given the uncertainties linked to the isotopic enrichment
factor between enamel and diet and to the simplified model using
the C3 browser/C4 grazer dichotomy, estimations of paleodiets
must be treated with great caution. Although global patterns are
interpretable in terms of differences between C4 grazers and C3
plant eaters, it is more difficult to conclude about the nature of
the consumed C3 plants.
Diagenesis is one of the biggest potential problems when
applying stable isotope biogeochemistry to fossils (Wang &
Cerling 1994). Numerous methodologies were recently devel-
oped to evaluate, or even correct, the effect of diagenesis on
isotopic compositions of fossilized enamel, but they are rarely
applied (e.g. Zazzo et al. 2004).
33
 Part I: Evolution, Taxonomy, and Domestication

Results and Discussion of the third molars, and retains strongly developed premolars
up to its extinction at the end of the Pleistocene (Gilbert 2008;
Figures 3.1, 3.2, and 3.3 illustrate the morphological changes in
Souron et al. 2015a). Kolpochoerus majus shares some cranio-
third molars in the Nyanzachoerus–Notochoerus, Kolpochoerus–
mandibular and dental features with the extant Hylochoerus but
Hylochoerus, and Metridiochoerus–Phacochoerus groups,
transitional forms are unknown, the biggest morphological dif-
respectively, compared to the changes in stable carbon isotopes
ference being the strong reduction of incisors and premolars in
of their enamel.
Hylochoerus (Souron et al. 2015a).
Most species of Nyanzachoerus display cranio-mandibular
δ13C values of Kolpochoerus older than 4 Ma indicate mixed
and dental morphologies typical of omnivorous suids. Their
diets of C3 and C4 plants. As early as 3.8 Ma for some specimens
third molars align well with the omnivorous morphotypes
of K. afarensis, and after 3 Ma for most specimens, the inferred
described earlier (Figure 3.1). Those species with omnivorous
diets are strongly dominated by C4 plants, even for the den-
morphotypes also display much enlarged third and fourth pre-
tally more conservative K. majus. Some populations of derived
molars with a morphology convergent to that of bone-crushing
Kolpochoerus from South Africa were consuming mostly C3
hyaenids. It has therefore been proposed that Nyanzachoerus
plants, but those could have been C3 grasses as all the plants
species were omnivorous but with a specialization to process
there were C3 plants (Figure 3.2; Luyt et al. 2000). Note that the
hard objects such as bones or nuts (e.g. Cooke and Wilkinson
C3 plants-dominated diet in extant Hylochoerus does not mean
1978). Nyanzachoerus jaegeri documents the transition with
it is a browser (contra Harris & Cerling 2002; Cerling et al. 2015)
Notochoerus around 4 Ma (this species is often placed in the latter
as the specimens analysed come from high-altitude areas where
genus Notochoerus) with more elongated third molars. Species of
grasses mostly use the C3 photosynthetic pathway. Other stud-
Notochoerus display cranio-mandibular and dental adaptations
ies indicate a mixed diet of various kinds of plants (including
to herbivory. They are extremely derived dentally with third
grasses) in the extant giant forest hogs (e.g. d’Huart 1978).
molars that are very elongated and very high-crowned (Figure
Early members of Metridiochoerus from 3.8 Ma to 2.5 Ma
3.1). They also display reduced premolars and incisors, roughly
are characterized by cranio-mandibular and dental morpholo-
similar to those of the extant Hylochoerus and Phacochoerus
gies typical of omnivorous suids. Starting around 2.5 Ma, all the
(Harris & White 1979). Those morphological changes are asso-
species display third molars that align with herbivorous mor-
ciated with similarly dramatic increases in body size.
photypes (Figure 3.3). There is a dramatic increase in length and
Stable carbon isotope values of early Nyanzachoerus indicate
height of third molars in the lineage M. andrewsi followed by
diets dominated by C3 plants up to around 5 Ma (Figure 3.1). After
an even more derived M. compactus. There is another group of
5 Ma, most Nyanzachoerus and Notochoerus display δ13C values
Metridiochoerus with third molar morphologies very close to
indicating diets dominated by C4 plants (Figure 3.1). However,
that of Phacochoerus.
note that the proportion of C3 plants consumed remains non-
In Metridiochoerus, δ13C values as early as 3 Ma indicate diets
negligible and even important in some cases. For example,
strongly dominated by C4 plants. Again, a few South African
in Plio–Pleistocene sites of the Malawi Rift, the paleodiets of
specimens and extant warthogs from high-altitude areas dis-
Nyanzachoerus and Notochoerus specimens were strongly domi-
play diets with strong C3 signals (Figure 3.3), but that does not
nated by C3 plants (Lüdecke et al. 2016). The authors interpreted
mean they were not consuming C3 grasses in abundance.
those data as suggesting a more closed habitat than in con-
A similar pattern is observed in Kolpochoerus and
temporaneous sites of eastern Africa, but an equally plausible
Metridiochoerus with strong discrepancies between the diets
hypothesis would be that those suids were consuming C3 grasses.
as expected from the dental morphology and the diets as
Overall, the δ13C values of the Nyanzachoerus–Notochoerus
reconstructed from the δ13C values. Pliocene specimens with
group are in good agreement with the ecomorphological recon-
third molars that would align with the omnivorous morpho-
structions, indicating omnivorous diets before 5 Ma and then a
types consumed large amounts of abrasive C4 plants, presum-
shift towards herbivory after 5 Ma. However, we cannot rule out
ably grasses. In both groups, the strong morphological changes
the hypothesis that specimens older than 5 Ma were consuming
(most notably the increase in length and height of third molars)
large amounts of C3 grasses or forbs.
that take place during the Pleistocene are not correlated with an
Species of Kolpochoerus from the first occurrence around
increased consumption of C4 plants. The key adaptations that
5.5 Ma to around 3 Ma are characterized by cranio-mandibular
allowed those extinct suids to enter the herbivorous niches were
and dental morphologies typical of omnivorous suids, nota-
therefore likely not the dramatic changes of the third molars.
bly with third molars very similar to those of Potamochoerus
(Figure 3.2). Around 3 Ma, there is a major cladogenetic split
with K. afarensis giving rise to two lineages. The first lineage,
Kolpochoerus limnetes (also called K. heseloni), displays a pro-
Towards a New Scenario of Adaptations to
gressive anagenetic trend towards more and more elongated Herbivory in African Suids
third molars and gave rise around 1.5 Ma to much more derived As already observed by Harris and Cerling (2002) on more lim-
species with more elongated and especially higher-crowned ited data, the review conducted here indicates that there is a
third molars (Figure 3.2). The premolars are reduced in the strong discrepancy between the changes in dental morphology
younger specimens. The second lineage, starting with K. phil­ and the changes in diets as indicated by stable carbon isotopes.
lipi then continued by K. majus, is more conservative dentally To understand the herbivorous adaptations of the African suids,
and displays a more moderate increase in length and height it therefore seems crucial to use a more integrative approach
34

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to place the morphology and diet of the suids within a larger
physiological (digestive system, body size, etc.) and ecological
context. When feeding, herbivorous mammals have evolved
different trade-offs between multiple morphological, physi-
ological, ecological, and behavioural variables to get enough
nutrients from the consumption and digestion of food: time
spent feeding, quantity of food ingested, efficiency in reducing
the food into digestible items, type and duration of digestion
(multi-chambered stomach, simple stomachs, rumination, cae-
cotrophy, etc.), body size. Those trade-offs have focused much
research and literature in ruminants and equids (e.g. Clauss
et al. 2008a) but no one, as far as the author knows, ever pro-
posed an explanation of suid herbivorous adaptations taking
into account those trade-offs. Here I propose a first preliminary
version of such a scenario.
First, previous studies considered both the length and height
of the third molars as comparable proxies for the degree of her-
bivorous adaptations. However, in terms of functional morphol-
ogy, both variables are obviously controlled by different factors.
Elongation of the third molars is likely related to chewing effi-
ciency as it multiplies the number of chewing sites. Elongated
teeth with numerous chewing sites are efficient for chewing
small food items (Lucas 2004: 169–170) and give an advantage
on a day-to-day fitness. Hypsodonty is rather linked to the dura-
bility of the tooth, i.e. its capacity to resist wear caused by phyto-
liths and grit contained in the food. A high crown gives a fitness
advantage on the scale of a lifetime.
The behaviour and ecology of extant warthogs within the
guilds of African herbivorous mammals are unique. Field obser-
vations indicate that they are able to feed on minute fragments
of grasses. Warthogs frequently feed on grass fields so short that
https://doi.org/10.1017/9781316941232.005 Published online by Cambridge University Press
Chapter 3: Diet and ecology of extant and fossil wild pigs
the grass blades are barely visible. I myself observed common
warthogs feeding on lands in the Pilanesberg National Park,
South Africa, shortly after fires: to a human eye, the land looked
pretty much barren, but the warthogs were able to feed on tiny
blades of grasses, and they were the only ungulates feeding there
(Figure 3.4). Similarly, warthogs are able to feed on grasslands
after they were grazed very short by other ungulates. By anal-
ogy with the extant warthogs, I propose here that most extinct
herbivorous suids that displayed dramatic adaptations of the
third molars were specialized on a diet of very small food items,
ingested in small mouthfuls.
Contrary to other ungulates (ruminants, tylopods, hippos,
tayassuids, equids, etc.), must suids do not display any special-
ized adaptations of the soft tissues of their digestive system to
cope with the plant matter, and especially the grasses. Their
stomach is composed of only one chamber, they do not rumi-
nate, and they do not have an enlarged caecum. That is also true
of the warthogs, even though their diets are composed almost
exclusively of grasses (Clauss et al. 2008b). Due to their limita-
tions in terms of digestive capacities of plant matter, herbivo-
rous suids need to reduce as much as possible the size of the
plant particles that enter the stomach.
A suite of morphological, ecological, and behavioural adap-
tations evolved to mitigate that problem: how to digest vegetal
matter without a specialized digestive physiology and stomach
anatomy? In suids, the emphasis when feeding on grass is the
selection of small and tender grass (ecology, behaviour) and the
buccal reduction of the grass into tiny digestible particles before
swallowing. On the contrary, equids and bovids are less limited
in terms of the types of vegetation they can feed on and the focus
is more on the post-swallowing digestion.
Figure 3.4  Picture of a common
warthog (Phacochoerus africanus)
feeding on near-barren soil in the
Pilanesberg National Park, South
Africa. Photo by Antoine Souron.
35
 Part I: Evolution, Taxonomy, and Domestication
The dramatic dental adaptations in African suids can there-
fore be considered as a solution to this particular problem: how
can small mouthfuls of small food items be processed efficiently
before digestion? The smaller the food item, and the smaller the
number of food items ingested at once (the size of the mouth-
ful), the less likely it is to break efficiently all the food items
with a small number of chewing cycles. The elongation of the
third molars with multiplication of cusps increase the length of
available enamel crests available for breaking small fragments
of grasses. A thin enamel also allows more numerous pillars to
be packed into the same dental surface, therefore increasing the
ratio of enamel crests to dental surface.
Herbivorous pigs display teeth that maximize the ratio of
enamel crests to the masticating surface. Multi-cuspidy, thin
enamel, and tightly packed cusps all enable a great ratio of
enamel crests length to dental surface to maximize the number
of breaking sites when masticating small elements of grass.
The specialization on small mouthfuls of short grass and the
maximization of the breaking sites enable herbivorous pigs to
break grass cells enough to digest their nutrients, which circum-
vents their lack of rumination and multi-chambered fermenting
stomach.
The height increase could represent an adaptation to the
abrasiveness of the grasses, because of its high content in exog-
enous mineral grit and silica phytoliths (Merceron et al. 2016).
Alternately, in the proposed scenario for suids, height increase
could also correspond to a strong attrition (wear caused by con-
tact between the upper and lower teeth during mastication) due
to the specialization on small mouthfuls of small food items.
The C3 component of the herbaceous stratum during the
Neogene and Quaternary is poorly known so far. If the afore-
mentioned scenario is confirmed, and extinct suids did have a
preference for tender, short grasses and forbs, they were maybe
highly selecting C3 herbaceous plants in the landscapes. In the
Malawi Rift, for example, extinct suids fed extensively on C3
plants whereas C4 grasses were present (as evidenced from their
consumption by some bovids, Lüdecke et al. 2016). Therefore,
better documenting and understanding the paleoecology of
herbivorous suids (and other herbivorous mammals with lim-
ited digestive capacities, such as cercopithecid primates of the
genus Theropithecus) could have strong implications for the
paleobiology of our own relatives, who also evolved adaptations
to herbivory (in Paranthropus) in the same landscapes.
Perspectives
In the future, it will be crucial to go further than the current
analyses that are mostly limited to stable carbon isotopes and/or
basic quantification of morphology. Several methodologies will
prove extremely useful when combined with the existing data.
Additional biogeochemical proxies for paleoecology are
promising. They will be all the more informative if they are
applied to serial samples documenting the intratooth variations
corresponding to the seasonal variations in diet and environ-
ment. For example, stable oxygen isotopes (δ18O) are related
to the degree of water dependence of the animal and also the
local rainfall amount in tropical Africa (Harris & Cerling,
2002; Souron et al. 2012). Some major elements may be related
36
https://doi.org/10.1017/9781316941232.005 Published online by Cambridge University Press
to the consumption of underground storage organs of plants
(Sponheimer et al. 2005) and stable magnesium isotopes dis-
tinguish well omnivorous mammals from herbivorous ones
(Martin et al. 2015). Dental microwear, the study of the micro-
scopic scars left at the surface of the enamel facets during the
mastication of food items, discriminates fine differences in diets
(e.g. Merceron et al. 2016 for a controlled feeding experiment on
sheep Ovis aries with diets of grasses versus forbs). Recently, 3D
dental microwear texture analyses were developed and provide a
quantified way to compare the microwear features across mam-
mals (see the recent review by Calandra & Merceron, 2016).
Souron et al. (2015b) notably discriminated among extant suids
between omnivorous, mixed feeders, and grass-eating species.
Ecomorphology of postcranials, a methodology already
applied to African extant and extinct suids, could also provide
additional insights into their ecology and habitat (Bishop 1999).
The main limitation is the paucity of fossilized postcranial
remains associated with cranio-mandibular or dental speci-
mens so that they can be assigned with confidence to a taxon.
Morphological adaptations to new diets also take place at
the microscopic scale of the components of the mammalian
enamel. Microscopic bioapatite prisms are arranged in dif-
ferent ways depending on the phylogeny and ecology of the
mammals according to the biomechanic constraints sustained
during mastication. For example, the premolars of bone-
crushing hyaenids display peculiar enamel microstructures
related to their diet (Tseng 2012) and could be compared to the
premolars of Nyanzachoerus to test their function. Documenting
the changes in enamel microstructure in African suids, along
the transition from omnivory to herbivory, could explain how
extinct suids with omnivorous morphotypes (such as early
members of Pliocene Kolpochoerus and Metridiochoerus) were
able to consume large amounts of abrasive C4 grasses.
Overall, the future of suid paleoecology is bright. Integrating
numerous complementary and independent methodologies
will without doubt allow us to better understand the timing
and rhythm of herbivorous adaptations, and the relationships
between diet, morphology, and environmental constraints.
That will allow many so far unanswered questions to be
addressed. How were the different sympatric species of herbivo-
rous suids sharing the niches? Was there competition for dietary
resources with other herbivorous mammals (ungulates, cerco-
pithecid and hominid primates, etc.) and omnivorous/carnivo-
rous mammals (e.g. hyaenids)? Which factors could explain the
lack of success of African herbivorous suids in Eurasia despite a
few limited attempts to disperse from Africa into Eurasia?
Acknowledgements
I thank the editors and Cambridge University Press for invit-
ing me to contribute to this volume, and especially Mario
Melletti for his infinite patience. The ideas developed in this
review stem from almost seven years of reflections on the evo-
lutionary history of our beloved Suidae. This brainstorming
was triggered and sustained by the constant support, mentor-
ing, and stimulating discussions of Jean-Renaud Boisserie and
Tim White, as well as numerous other colleagues. I am also
extremely grateful to Jean-Renaud Boisserie for permission

to use photographs of the fossils collected by the Omo Group
Research Expedition, and to Tim White for permission to
use photographs of the fossils collected by the Middle Awash
research project and of the casts of fossil suids from the collec-
tions of the Human Evolution Research Center, University of
California at Berkeley. I also thank John Rowan for improving
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