Evolving Earth 1 (2023) 100005

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Evolving Earth
journal homepage: www.sciencedirect.com/journal/evolving-earth

Ecological polarities of African Miocene apes

Gregory J. Retallack
Department of Earth Sciences, University of Oregon, Eugene, OR, 97403-1272, USA

A R T I C L E I N F O A B S T R A C T

Keywords: Humans have been considered ecologically unspecialized, and our evolution a compromise path through a maze
Miocene of conflicting influences. Generalist ecological roles contrast with ecological polarities such as competitor, breeder
Kenya or tolerator. Ecological polarities can be approximated in fossil mammals by relative size of canines for com-
Proconsul
petitors, incisors for breeders, and molars for tolerators. Considered in this way, early Miocene apes of Kenya were
Ekembo
Kenyapithecus
generalists, but show a greater range of ecological polarity than modern apes, including ecological polarities
Victoriapithecus found in living apes, macaques, and vervets. As many as six primates in a single Miocene aleosol show diverse
ecological polarities, implying competitive exclusion. Middle Miocene monkeys and apes were more molarized
and marginally more tolerant than early Miocene primates, so more like humans in that respect. This adaptive
shift of 15 Ma was at a time of climatic aridity and open vegetation indicated by associated paleosols. A 20 m.yr
record of Kenyan paleoprecipitation from paleosols indicates that 15 Ma was unusually dry after exceptionally wet
paleoclimate of 16 Ma. This new Kenyan paleosol record of paleoclimate is from the same localities as the fossil
apes, and uninfluenced by whole ocean mixing, salinity and ice volume effects which compromise similar marine
isotopic proxies of global change. Molarization of apes at 15 Ma ago was only one of a series of adaptations
selected by at least 9 dry alternating with wet episodes which shifted forest to shrubland ecotones over the past
20 m.yr. Our evolutionary lineage ran a gauntlet of Neogene climatic and vegetation changes in Africa by
adopting a generalist ecological role.

1. Introduction here is “ecological polarities”, for axes of relative proportional size of

1.1. Paleoecological question
Humans and other primates have long been considered ecologically
unspecialized (Simpson, 1949), and this study aims to quantitatively
evaluate that view. Human limbs retain a pentadactyl plan, rather than
elongated limbs of horses or the flippers of whales. Humans do not have
huge canines for threat or predation, like sabre-tooth cats and other
competitors. Nor are humans adapted to resource scarcity, like camels,
and other tolerators. Nor are humans as fecund as rodents, or other
breeders. Competitors, tolerators and breeders are three evolutionary
polarities, or roles toward which organisms may specialize (Retallack,
2004). These observations can be quantified by simple measurements
and plotting within a triangular diagram for proportional representation
of these extremes, with specialist species out toward the apices and
generalist near the middle (Fig. 1).
Competitor, tolerator and breeder ecological roles have been widely
quantified by plant ecologists as evolutionary “strategies” (Grime, 1979)
and “dimensions” (Westoby et al., 2002), but the terminology preferred
adaptive features (Retallack, 2004). The term polarity avoids anthropo-
morphism of “strategy” and simple linear connotation of “dimension.”
This is not the same as evolutionary polarity in the sense of relative
ordering of primitive or derived, apomorphic or plesiomorphic charac-
ters (Stevens, 1980; Bittrich and Struck, 1989), although Andews (1982)
has used “ecological polarity” in that sense of evolutionary polarity. Nor
is it the same as developmental or cell polarity, such as differentiation of
anterior or posterior (Kropf, 1992; Campanale et al., 2017). The aim of
ecological polarity studies is to quantify adaptation to particular niches.
Other animals may have been at a competitive, reproductive, or tolerance
disadvantage during extremes of global change, but our generalist line-
age may have persisted by virtue of flexibility to cope with spatial and
temporal disturbances at small population sizes (Potts, 1996).
This study quantifies and tests such ecological generalizations for
early and middle Miocene primates of Kenya (Fleagle and Kay, 1985;
McNulty et al., 2015; Alm
ecija et al., 2021), and a variety of modern
primates (Fleagle, 2013). These extinct apes are largely known from
fossil teeth. A previously published morphospace (Retallack, 2004)
chosen to approximate mammalian ecological polarities was the relative

E-mail address: gregr@uoregon.edu.

https://doi.org/10.1016/j.eve.2023.100005
Received 15 June 2023; Received in revised form 3 August 2023; Accepted 7 August 2023
2950-1172/© 2023 The Author. Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
G.J. Retallack
Fig. 1. Ecological polarity concept.
emphasis of functional areas of their three main tooth types, canines,
incisors and molars. Simple measures of upper tooth dimensions were
used to compute the functional area of molars, incisors and canines, and
the proportion of the sum of the three tooth areas for each tooth type
(Fig. 2). Primate canine size is significantly correlated with both intra-
specific and interspecific aggression (Kay et al., 1988; Manning and
Chamberlain, 1993; Kappeler, 1996). Mammalian incisor proportions
show significant correlation with fecundity measured as offspring per
year (Retallack, 2004). Mammalian molar occlusal area is strongly
correlated with body mass, home range size, basal metabolic rate, life
span, gestation time and age at first birth, and inversely correlated with
fecundity and population density (Martin, 1990; Janis and Carrano,
1992). The ecological polarity approach of ratios of areas factors out
body mass, so that relative molar size represents metabolic and popula-
tion characters of tolerator polarity (Retallack, 2004). Each species can
be plotted as a point with a standard deviation envelope within a trian-
gular diagram with apices of 100% canines, 100% molars and 100%
incisors (Fig. 1). Within this convention, rodents are more than 50%
Fig. 2. Measurements and formulae for calculating ecological polarity of mammals demonstrated on skull of Ekembo heseloni.
2
Evolving Earth 1 (2023) 100005
incisorized (lower left hand corner), sabre-tooth cats more than 50%
caninized (upper corner) and ungulates more than 50% molarized (lower
left hand corner).
While differences in incisorization, molarization and caninization are
significant across all mammals (Retallack, 2004), does that extend also to
the lesser variation in primates? Modern and fossil apes, monkeys and
lemurs occupy the central triangle of generalists (Retallack, 2004), in a
clear visualization of common wisdom (Simpson, 1949). The teeth of
primates are ecologically unspecialized compared with other mammals,
so this study quantifies limited primate variation in tooth proportions.
Incisorization in primates has been regarded as a “sclerocarp syndrome”
for feeding on hard shelled fruits (Kinzey, 1992; McCrossin and Benefit,
1997; Ward et al., 1997; Kay, 2010), which may parallel the toughness,
small size, and high nutritional value of seeds fostering fecundity in ro-
dents (Retallack, 2004). Molarization in primates has been linked to
declining encephalization and less complex behavior in primates, but not
without exceptions (Jim
enez-Arenas et al., 2014). Some primates have
molarized premolars (Selig and Silcox, 2022), but premolars were not
measured in the present work for consistency with prior work (Retallack,
2004). Caninization in primates is primarily linked with sexual dimor-
phism and agonistic behavior during mate isolation and selection
(Thor
en et al., 2006), but female canine size also increases with male
canine size (Plavcan et al., 1995; Plavcan, 1998). This can be a problem
for evaluating fossils depending on the proportion of male or female
canines measured, although sex has been interpreted for many fossil
specimens (Pickford, 1985; Kelley, 1986, 1995; Pickford et al., 1999).
1.2. Geological background
This study is limited to data from around Lake Victoria for the early-
middle Miocene Nyanza Rift and Lake Victoria Basin (Fig. 3), which has
long yielded important Miocene primate fossils (Leakey, 1952; Pickford,
1986a; Tuttle, 2006). These localities are on a subsidiary rift to the main
East African or Gregory Rift, and were characterized by terrestrial
carbonatite-nephelinite volcanism, with a few lacustrine horizons
(Pickford, 1986b, 1995; Retallack, 1991; Bestland and Krull, 1999;
Lukens et al., 2017; Michel et al., 2014, 2020; Peppe et al., 2023).
Particular paleosols were named by descriptive terms in local lan-
guages, and characterized chemically and petrographically by Retallack
(1991, Retallack et al., 1995, 2002). Choka and Tut paleosols at Songhor
G.J. Retallack Evolving Earth 1 (2023) 100005

Fig. 3. Geological map of the Nyanza Rift, southwest Kenya (adapted from Retallack, 1991).

and Kwar paleosols at Koru are all roughly equivalent in age, 20.3
0.5
Ma by corrected K–Ar method for bracketing tuffs (Supplementary In-
formation Table S1), and represent lowland and volcanic flank paleo-
environments respectively (Bishop et al., 1969; Retallack, 1991). On
Rusinga Island, Chido paleosols in the Kiahera Formation some 18 Ma in
age are stratigraphically lower than Tek paleosols in the Hiwegi For-
mation some 17.8
0.16 Ma by K–Ar (Drake et al., 1988; Retallack et al.,
1995; Bestland and Krull, 1999). Yom paleosols of a former alluvial
lowland (dambo) on Maboko Island are beneath a tuff dated by 40Ar/39Ar
at 14.71
0.16 Ma (Feibel and Brown, 1991; Retallack et al., 2002).
Chogo and Onuria paleosols at Fort Ternan formed on volcanic slopes at
about 13.7
0.3 Ma by 40Ar/39Ar (Bishop et al., 1969; Pickford et al.,
2006). By mid-late Miocene the subsidence and sedimentation in the
Nyanza Rift was waning (Pickford, 1986a, 1986b), but paleoenvir-
onmental time series of radiometrically dated paleosols can be extended
to the present in the Gregory Rift: in the Tugen Hills of central Kenya for
the mid-late Miocene (Jacobs and Deino, 1996; Deino et al., 1990;
Behrensmeyer et al., 2002; Pickford and Kunimatsu, 2005; Senut et al.,
2000), and the Turkana Basin of northern Kenya for the late Miocene to
Quaternary (Wynn, 2001, 2004a, 2004b).
2. Materials and methods
2.1. Taxonomic basis
Taxonomy of Kenyan Miocene apes used here generally follows
Harrison (2002, 2010), Ward and Duren (2002), Pickford and Kunimatsu
(2005), and Alm
ecija et al. (2021). Only those taxa that could be linked to
specific paleosols and sites (Pickford, 1986a; Retallack, 1991) were
included in this analysis (Table 1), other specimens from other sites or
subsequently described without precise locality data are not considered.
Victoriapithecus macinnesi is retained as a valid species (Benefit, 1987,
1993; Benefit and McCrossin, 1997; Miller et al., 2009), rather than
emended to the genus Prohylobates (Pickford and Kunimatsu, 2005).
Ugandapithecus major is accepted in place of Proconsul major (Le Gros
Clark and Leakey, 1951; Harrison, 2010), because its large postrcrania at
Songhor and in Ugandan localities are distinct from those known in
Proconsul (Senut, 2016, Senut et al., 2000; Gommery et al. (2002).
Ekembo nyanzae and E. heseloni are used here instead of Proconsul,
although that genus is retained for Proconsul africanus and P. meswae
(McNulty et al., 2015). Equatorius africanus is used here, rather than the
genus Kenyapithecus (following Ward et al., 1999; Kelley et al., 2002,

Table 1
Tooth measurements of selected Miocene taxa and localities from Kenya.
Taxon Locality Canines Incisors Molars Source

area (mm2) error no. area (mm2) error no. area (mm2) error no.

Dendropithecus macinnesi Hiwegi, Rusinga Is. 32.72 13.47 9 31.36 8.91 12 49.92 7.6 13 Andrews (1978)
Dendropithecus macinnesi Songhor 33.61 7.84 4 32.24 12.9 5 50.4 0 1 Andrews (1978)
Equatorius africanus Maboko 53.47 12.17 4 43.54 21.61 4 111.02 19.9 4 Pickford (1985)
Kenyapithecus wickeri Fort Ternan 78.23 5.2 2 56.31 29.85 2 119.4 15.1 3 Pickford (1985)
Limnopithecus legetet Songhor 17.89 3.11 4 27.46 5.04 8 35.48 4.16 18 Andrews (1978)
Proconsul africanus Koru 57.61 6.67 2 48.65 19.3 2 80.57 14.8 9 Andrews (1978)
Ekembo heseloni Kiahera, Rusinga Is. 42.05 12.5 5 54.24 15.5 10 82.4 14 32 Andrews (1978)
Ekembo nyanzae Hiwegi, Rusinga Is. 31.71 1.62 2 84.1 24.1 14 131.16 28.1 16 Andrews (1978)
Rangwapithecus gordoni Songhor 41.25 1 0 60.9 13.04 4 83.03 18.4 12 Andrews (1978)
Ugandapithecus major Koru 159.71 34 2 92.43 39.98 3 162.26 44.5 16 Andrews (1978)
Victoriapithecus macinnesi Maboko 17.64 4.79 159 35.27 17.31 130 48.46 9.42 339 Benefit (1987)

3
G.J. Retallack
2008; Harrison, 2010), or the Eurasian genus Griphopithecus, contrary to
Begun (2007), Benefit and McCrossin (2000) and Pickford and Kuni-
matsu, 2005). Less controversial are the taxonomy of Dendropithecus
(Andrews and Simons, 1977), Limnopithecus (Harrison, 2002, 2010; Cote
et al., 2016), and Rangwapithecus (Hill et al., 2013; Cote et al., 2014).
2.2. Dental measurements of fossil apes
Dental measurements and formulae used to approximate ecological
polarity are shown in Table 1 and Fig. 2, with full data in Supplementary
Information Table S2. Only those remains from single collections and
known paleosols that represent attritional assemblages were used
(Retallack, 1991). The measurements are for individual teeth, both iso-
lated and within complete mandibles or maxillae. There has been no
adjustment to relative representation of teeth for a particular dental
formula: only relative sizes of the three tooth categories. Paleosol loca-
tions of these specimens in the Kenyan National Museum were estab-
lished from museum records, and discussions and correspondence with
collectors (Pickford, 1986a, 1986b).
2.3. Paleosol evidence of paleoclimate and ecosystems
Ecological polarity analysis of paleosols followed methods described
elsewhere (Retallack, 2004), and included measurement of soil fertility
(100 times the molar ratio of bases, MgO, CaO, Na2O and K2O, to Al2O3),
of soil youth (100 minus soil development index out of 100) and of
hardship, such as aridity indicated by shallow depth of calcic horizon
(1000 divided by depth to Bk in cm).
Depth to carbonate horizon (D in cm) in soils is related to mean
annual precipitation (P in mm) by equation (1) below with R2 ¼ 0.52,
S.E. ¼
147 mm. p ¼ 0.00001 (Retallack, 2005). This depth can be
corrected for compaction due to overlying sediment using geological
estimates of overburden and standard formulae (Sheldon and Retallack,
2001). Also related to mean annual precipitation (P) is nutrient base
content (C–– Al2O3/(Al2O3þCaO þ MgO þ Na2O) in moles) of soil Bt
horizons by equation (2) below with R2 ¼ 0.72, S.E. ¼
182 mm, p ¼
0.00001 (Sheldon et al., 2002).
P ¼ 137.24 þ 6.45D – 0.013D2 equation 1
0.197C
P ¼ 221.12.e equation 2
Relevant Bk and geochemical data extracted from previous publica-
tions is tabulated in Supplementary Information Table S1, Kenyan
Cenozoic paleosol paleoclimatic data. A compilation of compaction cor-
rected depth to Bk in paleosols as a paleoprecipitation proxy is from
Retallack (1991, 2013), Wynn (2001, 2004a, 2004b), Wynn and Retal-
lack (2001) and Retallack et al. (2002). Relevant Kenyan paleosol
chemical data is from Retallack (1991), Retallack et al. (1995, 2002), and
Wynn and Retallack (2001). None of the Kenyan paleosol samples was
pretreated to remove carbonate as recommended by Michel et al. (2014),
so only samples with less than 5% CaO were used to calculate
paleoprecipitation.
Chemical weathering also alters the mineral content of soils, espe-
cially their clay minerals, which begin as smectites, and then lose cationic
bases with further chemical weathering to become kaolinite. This indi-
cation of paleoprecipitation works best with non-calcareous soils, which
are found in climates receiving more than 1000 mm mean annual pre-
cipitation (Sheldon et al., 2002). In Africa today, smectite is dominant in
soils receiving less than 1200 mm mean annual precipitation, and
kaolinite dominant in wetter climates (Mizota et al., 1988; Dworkin,
2023). Thus, non-calcareous smectitic soils also define a limited paleo-
climatic window of between 1300 and 1000 mm mean annual precipi-
tation, which can be inferred from clay mineral data of Retallack (1991)
and Behrensmeyer et al. (2002).
Paleoprecipitation estimated from foliar physiognomy of three leaf
fossil localities, Kabarsero (12.6 Ma), Waril (8.6 Ma) and Kapturo (6.8
4
Evolving Earth 1 (2023) 100005
Ma) by Jacobs (200a, b) represent unusually humid conditions for the
region (Supplementary Information Table S1).
3. Results
3.1. Ecological polarities of Miocene apes and monkeys
Ecological polarity analysis gives insight into the puzzling diversity of
Kenyan Miocene apes. There are only eight living species of great apes
(Hominidae) today, with largely disjunct ranges (Fleagle, 2013), yet
single paleosols such as those at the 20 Ma fossil site of Songhor contain
as many as 6 species of sympatric stem-apes (Retallack, 1991). These
assemblages are attritional accumulations of bones broken and weath-
ered during soil formation (Pickford, 1986b), and so represent former
communities, rather than regionally homogenized assemblages found in
lake or stream deposits (Verhaegen et al., 2002). Their paleosols also
show little evidence of gleization due to waterlogging (Retallack, 1991;
Retallack et al., 2002), and no support for “aquarboreal human ances-
tors” (Verhaegen et al., 2002).
Each ape species plots in different regions of the generalist central
triangle of incisor-canine-molar morphospace (Fig. 4). Ugandapithecus
major, for example, shows greater caninization than the other species and
is most similar in that respect to living orangutans (Pongo pygmaeus). The
large size and limbs of Ugandapithecus major appear adapted for slow,
suspensory climbing (Gommery et al., 2002; Senut, 2016), again similar
to orangutans. Other taxa (Dendropithecus macinnesi, Proconsul africanus)
have caninization more like living macaques, and yet other taxa (Lim-
nopithecus legetet, Rangwapithecus gordoni) are more like living vervets in
this respect. These pairs may be further divided by their postcranial
anatomy, which indicate that Dendropithecus macinnesi was an under-
branch suspensory feeder, whereas Proconsul africanus, and Rangwapi-
thecus gordoni were overbranch climbers and feeders (Andrews and
Simons, 1977; Walker and Shipman, 2005; Harrison, 2010; Fleagle,
2013). Limnopithecus may also have been suspensory, but its postcrania
are not unambiguously known (Harrison, 2010). Ecological polarity
analysis thus confirms that Miocene apes were not only similar to modern
great apes, but to modern monkeys as well (Andrews, 1996; Alm
ecija
et al., 2021). The taxonomic diversity of Kenyan Miocene apes was
matched by their ecological diversity, exploiting a variety of forest tiers,
branch sizes, fruits, and leaves.
Modern Homo sapiens straddles the 50% molarization line like Ekembo
heseloni of 18 Ma on Rusinga Island, and both the ape Kenyapithecus
africanus and early monkey Victoriapithecus macinnesi of 15 Ma on
Maboko Island (Fig. 4). In contrast, generalist and apelike adaptations are
occupied by Ekembo nyanzae and Dendropithecus macinnesi of 18 Ma on
Rusinga Island and Kenyapithecus wickeri of 13.7 Ma at Fort Ternan. The
morphospace occupied by Dendropithecus macinnesi was unchanged from
20 to 18 Ma, but change in other lineages toward the tolerator pole
implies environmental hardship at 18 Ma and again at 15 Ma. Tough-
coated fruits are indicated by large procumbent incisors in 18 Ma
Ekembo nyanzae and 15 Ma Equatorius africanus, as well as Turkanapi-
thecus and Afropithecus from 18 Ma deposits in northern Kenya (Leakey
et al., 1988a, 1988b; Boschetto et al., 1992; McCrossin and Benefit, 1997;
Leakey and Walker, 1997; McNulty et al., 2015). Hard or fibrous foods
also are indicated by thick enamel and procumbent incisors of Equatorius
africanus and K. wickeri (Martin, 1985). Tough coated fruits and gourds
are also indicated by 18 Ma seeds of Annonaspermum (custard apple) and
Lagenaria (calabash) on Rusinga Island (Collinson et al. (2009) and 13.7
Ma seeds of Annonaspermum (custard apple) at Fort Ternan (Retallack,
1992). Furthermore, both Equatorius africanus and Victoriapithecus mac-
innesi have postcranial anatomy indicative of facility for ground-walking
like that of macaques and vervets, respectively (McCrossin et al., 1998).
These adaptations may have been selected by open and fibrous vegeta-
tion at 18 and 15 Ma, unlike more mesic vegetation of 20 and 16 Ma.
G.J. Retallack Evolving Earth 1 (2023) 100005

Fig. 4. Ecological polarity of selected species of Miocene apes from Kenya (C–G) and of their Miocene paleosols (A), as well as of comparable modern primates (B).
Envelopes around the point for each species represents the standard deviation of that measure. These measurements are not for complete specimens but for known
teeth. A generalist ecological role is indicated for fossil and modern primates, but deviations toward molarization are accompanied by evidence of aridity in asso-
ciated paleosols.

5
G.J. Retallack
3.2. Ecological polarity of paleosols
Confirmation of environmental changes at these times comes from
paleosols which entomb these fossil primates. The ecological polarity of
soils and paleosols can be approximated by three measures: (1) soil
fertility (molar ratio of bases to alumina), (2) soil youth (inverse soil
development index) and (3) soil hardship such as aridity (shallow depth
of soil carbonates). Competitor plants and animals are on the most fertile
soils, breeder plants and animals on youthful disturbed soils, and toler-
ator plants and animals on soils presenting hardships of drought or low
nutrition (Retallack, 2004). Weakly developed soils have bedding planes
and other original structures little disturbed by roots and weathering, but
as time progresses powdery carbonate accumulates and coalesces into
nodules of increasing size (Retallack, 2005) in a quantitative scale of
development which serves as a breeder polarity proxy (Retallack, 2004).
Tolerator polarity in calcareous soils can be inferred from depth within
paleosol profiles to carbonate nodules, which correlates significantly
with mean annual precipitation (Retallack, 2005), but needs to be cor-
rected for burial compaction in paleosols (Sheldon and Retallack, 2001).
Competitor polarity can be quantified by measures of chemical soil
fertility, such as bases/alumina ratios of non-calcareous parts of the soil.
The four major soil bases are important nutrients: Ca2þ for plant cell
walls and animal bones, Mg2þ central to chlorophyll; and Naþ and Kþ as
cytoplasmic electrolytes (Retallack, 2019). Such information from pale-
osols with fossil primates reveals selection pressures on their evolution.
There was clear ecological separation of some species to different
paleosols, but still high diversity within some paleosols (Fig. 5). In the 20
Ma sites of Koru and Songhor, Ugandapithecus shows little habitat spec-
ificity through a variety of tropical dry forest paleosols, but small taxa
(Kalepithecus, Micropithecus) are in upland soils and larger taxa (Proconsul,
Rangwapithecus, Dendropithecus, Nyanzapithecus) remained in lowland
forests close to water. One paleosol type (early successional Kiewo
pedotype: Fig. 5) has six taxa: including likely suspensory feeders (Lim-
nopithecus, Dendropithecus, Ugandapithecus, from small to large) and likely
overbranch feeders (Nyanzapithecus, Proconsul, Rangwapithecus, from
small to large). Thus there was niche partitioning of forest canopy tiers.
Diverse sympatric catarrhine communities persisted into the drier
woodland landscapes of Rusinga Island at 18 Ma, when paleosols with
the crumb peds and iron–manganese nodules of dambo grasslands (Yom
pedotype: Fig. 5) appear, but are rare and barren of primate fossils
(Retallack et al., 1995; Peppe et al., 2023). Other evidence for grasslands
of about the same age are abundant bunch grasses at Ugandan fossil sites
of Bukwa (Pickford, 2002) and Moroto (MacLatchy et al., 2023). Yom
paleosols of dambo grassland are common by 15 Ma on Maboko Island
(Retallack et al., 2002), and contain abundant vervet-like monkeys
(Victoriapithecus: note change of scale for this exceptional collection in
Fig. 5). Seasonally inundated grasslands of dry climates were not
encouraging to fossil apes (Lukens et al., 2017), which were more com-
mon in riparian woodland paleosols (Nyanzapithecus, Limnopithecus,
Mabokopithecus and Simiolus of Dhero pedotype). In contrast, Equatorius
at Maboko and Majiwa was wide ranging in both grassland (Yom, Dhero)
and pori woodland (Ratong), which it exploited more effectively than
other apes because of its thick enamelled, large molars useful for tough
foods (Martin, 1985), and its macaque-like limbs and feet (McCrossin
et al., 1998).
A similar pattern of forest dependence for Kenyapithecus and other
apes (Simiolus, “Proconsul”, nyanzapithecine: Harrison, 1992) persisted
within grassland woodland mosaics of Fort Ternan and Kapsibor at 13.7
Ma, when well-drained short-grass, wooded grassland was widespread
(Kappelman, 1991; Retallack, 1992; Dugas and Retallack, 1993). The
appearance of grasslands so encouraging for victoriapithecine ancestors
of vervets, was not so encouraging to apes, which remained rare com-
ponents of the fossil fauna.
Analysis of fossiliferous paleosols at these Kenyan sites demonstrate
that molarization increases in apes and monkeys corresponds with times
of shallow soil carbonate (Fig. 5), which indicates a paleoclimatic shift
6
Evolving Earth 1 (2023) 100005
from subhumid to semiarid climate (Fig. 4). Miocene apes and monkeys
adapted to aridification of their ecosystems and soils in different ways,
with apes remaining within woodland soils, but some monkeys abundant
within grassland soils.
4. Discussion
4.1. Extended paleoenvironmental time series for Africa
The time series of early to middle Miocene paleosols from the Nyanza
Rift and Victoria Lake Basin can be extended with comparable studies in
the mid-late Miocene Tugen Hills (Jacobs and Deino, 1996; Deino et al.,
1990; Behrensmeyer et al., 2002; Pickford and Kunimatsu, 2005; Senut
et al., 2001), and the Miocene to Quaternary Turkana Basin (Wynn, 2001,
2004a, 2004b; Liutkus-Pierce et al., 2019). This extended compilation
(Fig. 6) includes data on Kenyan paleosol depth to carbonate, chemical
and clay mineral composition. Depth to carbonate (Bk) increases with
mean annual precipitation in soils (Retallack, 2005), and so does the
degree of depletion of alkali and alkaline earth elements in subsurface
clayey (Bt) horizons of soils (Sheldon et al., 2002). Precipitation also
controls clay minerals of soil Bt horizons: smectite in dry regions (less
than 1200 mm mean annual precipitation) and kaolinite in wetter re-
gions (Mizota et al., 1988; Dworkin, 2023). Also included in this paleo-
climatic compilation (Fig. 6) are published inferences from size, shape
and features such as drip tips of fossil leaves (Retallack, 1992; Dugas and
Retallack, 1993; Jacobs and Deino, 1996; Jacobs, 2002a, 2002b; Michel
et al., 2014, 2020), which agree with estimates from paleosols where
both are found in close proximity. Non-calcareous paleosol mineral
composition and fossil leaf physiognomy both define isolated peaks in
paleoprecipitation, compared with the runs of numerous calcareous
paleosols (Fig. 6). Some of the volatility in this compilation represents
lateral landscape heterogeneity (Fig. 5), revealed in more detail by recent
studies (Michel et al., 2020; Peppe et al., 2023; MacLatchy et al., 2023).
Vegetation can also be reconstructed from known precipitation
ranges of modern African ecosystems (Anhuf et al., 1999; Sankaran et al.,
2005). This assumption fails for newly evolved ecosystems such as
grasslands, which can be detected in paleosols from fine root traces and
crumb ped structure, as well as from associated grass leaves, pollen, and
phytoliths (Retallack, 1992; Dugas and Retallack, 1993; Peppe et al.,
2023). The changing paleoclimatic range of grasslands has been recon-
structed from climatically sensitive features such as depth to Bk in the
same paleosols (Fig. 6). Grasslands and their soils (Mollisols) of uplands
(veld) and lowlands (dambo) were a newly coevolved ecosystem engi-
neered by grasses and grazers (Retallack, 2013). The expanded climatic
range of mollic paleosols of grasslands through time (Retallack, 2013) is
revealed by filled symbols in Fig. 6B. In contrast early successional and
forest communities had considerable antiquity in Africa, especially a dry
woodland community known as pori which was largely displaced by
grasslands in regions between 300 and 500 mm of mean annual precip-
itation (Retallack, 2007, 2012).
The extended paleoclimatic compilation reveals not just one Neogene
aridification and grassland expansion at about 7 Ma, as has long been
implied by the Miocene pluvial (Leakey, 1952), lake (Kent, 1944), and
rain forest (Andrews, 1996) hypotheses, and late Miocene grassland
hypothesis (Cerling, 1992; Cerling et al., 1997). These theories had
already been discredited by discovery of Miocene desert dunes, alkaline
lakes, grasses, grazing mammals, and grassland paleosols in East Africa
(Pickford, 1986b, 1995, 2002; Retallack, 1991; Retallack et al., 2002;
Lukens et al., 2017; Peppe et al., 2023; MacLatchy et al., 2023). Instead
the new data (Figs. 5–6) reveal a mosaic of habitats and a Neogene pa-
leoclimatic roller coaster of at least 9 desertifications, of which those at
18 and 15 Ma were among the driest. Of the intervening wet forests,
humidity spikes of 16 and 13 Ma were among the wettest.
The relationship between Miocene and living African apes are un-
clear, because of the extremely sparse fossil record of chimpanzees
(McBrearty and Jablonski, 2005) and gorillas (Suwa et al., 2007). The
G.J. Retallack Evolving Earth 1 (2023) 100005

Fig. 5. Paleosols of Miocene primates from southwestern Kenya [data from Retallack (1991) and Pickford (1986a, b), with taxonomy after Harrison (2002, 2010),
Retallack et al. (2002), Ward and Duren (2002), Alm
ecija et al. (2021), for selected stratigraphic levels.

7
G.J. Retallack
varied Miocene apes of southwest Kenya (Harrison, 2010) may have been
ancestral to erect-walking hominid fossils, such as Sahelanthropus (Brunet
et al., 2002), Orrorin (Senut et al., 2001), and Ardipithecus (White et al.,
2009; Lovejoy et al., 2009), which were still in woodland and forest
Alfisols from 6.5 to 4.2 Ma (Retallack, 2007). Starting with Austral-
opithecus anamensis our lineage sometimes ventured out onto grassland
Mollisols (Wynn, 2001; White et al., 2006), that had long supported an
adaptive radiation of baboons and vervets (Frost, 2007). Subsequent
8
Evolving Earth 1 (2023) 100005
Fig. 6. Compilation of paleosol data for 20 million
years of vegetation (A) and precipitation (B) variation
in Kenya, compared with carbon (C) and oxygen (D)
isotopic composition of marine foraminifera (Zachos
et al., 2001). Paleoprecipitation data from non-mollic
paleosols (B) is from depth to carbonate (open cir-
cles), clay mineralogy (diamonds), and chemical
composition (squares). Closed circles are those mollic
paleosols with carbonate as well as crumb peds and
fine root traces of grassland soils. Also included are
inferences from fossil floras (open triangles: Jacobs,
2002a, 2002b, Jacobs and Deino, 1996; Collinson
et al., 2009; Michel et al., 2014, 2020) and clay
mineral content (Retallack, 1991; Behrensmeyer
et al., 2002). Precipitation ranges of extant vegetation
types are from Holocene reconstructions (Anhuf et al.,
1999; Sankaran et al., 2005), and expansion of
grasslands from trace fossil and structural studies of
paleosols (Retallack, 2013).
australopithecines and humans of our lineage (Alm
ecija et al., 2021)
exploited C4 and C3 grassland mosaics of Mollisols, Alfisols, and Incep-
tisols after 4.2 Ma (Wynn, 2004a, 2004b; Wynn et al., 2006).
4.2. Local or global paleoclimatic change
The extended paleoclimatic compilation is similar to the oxygen
isotopic record from the deep-sea (Zachos et al., 2001), which has been
G.J. Retallack
used as a paleoclimatic proxy for evaluating human evolution in Africa
(de Menocal, 2004), but the match is not precise (Fig. 6B and C). A trend
of extreme and volatile middle Miocene values for paleoprecipitation and
marine stable isotopic composition, but subdued late Miocene to Qua-
ternary values is evident. The paleosol record reveals much greater
variation in rainfall than would be inferred from carbon isotopic
composition of marine foraminifera (Fig. 6C), which is damped by global
oceanic mixing with time lags of several thousand years (Zachos et al.,
2001). More profound damping is seen in oxygen isotopic composition of
marine foraminifera, which show a long-term increase in δ13C (‰) unlike
local rainfall and foraminiferal carbon records. This increase is plotted on
reversed axes in Fig. 6D because it has been interpreted as a record of
long-term temperature decline (Zachos et al., 2001; Veizer et al., 2001),
but is partly an artefact of long-term water recycling due to plate tec-
tonics (Kasting et al., 2006). The global oxygen isotope record also shows
an increase after 3 Ma due to continental ice cap sequestration of isoto-
pically light oxygen, in addition to temperature effects (Zachos et al.,
2001). The African paleosol record and global isotopic records thus
presents a concept of climatic variation experienced by our distant an-
cestors very different from past ideas of a seminal late Miocene climatic
event. Instead of a single origin of humanity at a turning point of envi-
ronmental change, the new record implies rather that our lineage
responded to a gauntlet of changing conditions with a variety of
adaptations.
Each fluctuation in climate and vegetation presented new crises and
opportunities to African hominoids. Some specialized features such as
procumbent incisors evolved by about 18 Ma in Afropithecus, Ekembo and
Kenyapithecus for hard-skinned fruits of dry woodlands (McCrossin and
Benefit, 1997). Several features of the femur in Ugandapithecus by 20 Ma
were adapted for climbing in woodlands (Senut, 2016). Yet other likely
forest adaptations such as erect stance in Orrorin and Sahelanthropus by 6
Ma (Brunet et al., 2002; Senut et al., 2000), and flat face in Kenyanthropus
by 3.5 Ma (Leakey et al., 2001) proved advantageous in the long term,
just as did likely aridland adaptations such as thick enamel in Afropithecus
and Equatorius by 18 Ma (Martin, 1985; McCrossin and Benefit, 1997;
Smith et al., 2003; Rossie and MacLatchy, 2013), adducted hallux in
Kenyapithecus by 15 Ma (McCrossin et al., 1998), and long legs and big
brains in Homo by 2.5 Ma (Bramble and Liebermann, 2004). Although
each of these ideas could, and should, be debated individually, the
general concept of human evolution as a generalist path through a
gauntlet of environmental challenges (Potts, 1996) is now supported by
the African record of fossil apes and soils.
5. Conclusions
Fossil vertebrates in attritional assemblages within paleosols are not
only a source of paleoecological information from adaptive features of
their skeletons, but selection pressures on those adaptations can be
inferred from the paleosols as well. Furthermore, paleosol diversity re-
veals habitat heterogeneity throughout the evolutionary history of our
apelike ancestors in Africa. Both fossils and paleosols can be considered
from the perspective of ecological polarities, which assess the relative
emphasis on competition, fecundity, and withstanding. Similarly, soils
can be ranked for their relative fertility, physical disturbance, and
hardship. Miocene paleosols and primates of southwest Kenya consid-
ered from this perspective showed increased molarization with dimin-
ished paleoprecipitation. As many as 6 sympatric species of apes in single
paleosols showed competitive exclusion in polarity ecospace. When
grasslands first appeared during the middle Miocene, early monkeys such
as Victoriapithecus, adapted to them just as vervets and baboons live in
grasslands today. However, early hominoid apes, such as Kenyapithecus,
remained in woodlands and forests. Grasslands played little role in evo-
lution of upright stance and other human adaptations until the Pliocene
appearance of australopithecines, whereas vervets and baboons had a
long history of adaptation to grasslands back to middle Miocene.
9
Evolving Earth 1 (2023) 100005
Declaration of interest statement
The author declares no conflict of interest in publishing this research.
Data availability
Data will be made available on request.
Acknowledgements
Fieldwork in 1984 and 1987 was undertaken with a permission letter
from Ms M.A. Mwango of the Office of the President of Kenya, and R.E.
Leakey of the Kenyan National Museum. Funding came from Wenner
Gren Foundation, and US NSF grants EAR8206183 and EAR8503232. S.
Gitau, H. Vallianatos, J. Wynn, E. Bestland, D. Dugas, G. Thackray and D.
Retallack aided fieldwork. Invaluable discussions and advice came from
P. Shipman, M. Pickford, B. Benefit and M. McCrossin.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.eve.2023.100005.
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