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Comparative dental development and microstructure of ... - UCL

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164 A. D. BEYNON ET AL.<br />

Beard et al., 1986; Begun et al., 1993;<br />

Fleagle, 1983; Harrison, 1987, 1993; Lewis,<br />

1971; Napier & Davis, 1959; Rose, 1997;<br />

Ward et al., 1991, 1993). Overall, these<br />

postcranial characters suggest Proconsul was<br />

an arboreal quadruped with a varied positional<br />

repertoire that indulged in relatively<br />

slow climbing but which showed few signs <strong>of</strong><br />

forelimb suspensory behaviour (Walker,<br />

1997). Evidence from the skull <strong>and</strong> dentition<br />

includes characters that link Proconsul<br />

with later hominoids. The estimated degree<br />

<strong>of</strong> encephalization <strong>of</strong> P. heseloni, although<br />

based on one specimen (KNM-RU 7290),<br />

suggests that Proconsul had a bigger brain<br />

than modern cercopithecoids <strong>of</strong> a comparable<br />

body mass (Walker et al., 1983). Other<br />

cranio<strong>dental</strong> features such as the presence <strong>of</strong><br />

a frontal air sinus (Walker & Teaford,<br />

1989), a wide frontal bone at bregma, the<br />

<strong>development</strong> <strong>of</strong> a maxillary jugum, a low<br />

crowned P 3 <strong>and</strong> reduced cusp heteromorphy<br />

<strong>of</strong> the upper premolars are also each considered<br />

by some to be hominoid synapomorphies<br />

(Andrews, 1985). For a recent<br />

review see Walker (1997).<br />

Four species <strong>of</strong> Proconsul are now<br />

described (Walker et al., 1993; Teaford<br />

et al., 1993; Andrews, 1996). Proconsul<br />

africanus <strong>and</strong> Proconsul major are known<br />

from the type sites <strong>of</strong> Koru <strong>and</strong> Songhor in<br />

western Kenya <strong>and</strong> P. major also from<br />

Meswa Bridge in Kenya <strong>and</strong> Napak in<br />

Ug<strong>and</strong>a. P. heseloni <strong>and</strong> P. nyanzae are the<br />

two species that are represented at Rusinga<br />

<strong>and</strong> Mfangano Isl<strong>and</strong>s in Kenya (Walker<br />

et al., 1993). Ruff et al. (1989) used crosssectional<br />

measurements <strong>of</strong> the femoral<br />

diaphysis <strong>and</strong> articular dimensions to estimate<br />

the body weight <strong>of</strong> Proconsul specimens<br />

from Rusinga <strong>and</strong> Mfangano. Rafferty et al.<br />

(1995) subsequently made estimates from<br />

ankle joint surface areas. Both estimates are<br />

around 9–12 kg for the smaller P. heseloni<br />

specimens (about the same as a siamang, or<br />

twice that <strong>of</strong> smaller Hylobates species).<br />

Body weight estimates for P. nyanzae are<br />

closer to those <strong>of</strong> female chimpanzees <strong>of</strong><br />

the smallest subspecies averaging 35·6 kg<br />

(Rafferty et al., 1995).<br />

Reviewing the paleoecology <strong>and</strong> the<br />

hominoid paleoenvironments, Andrews<br />

(1996) presented evidence that Songhor <strong>and</strong><br />

Koru (dated at 19–20 Ma) have fossil faunas<br />

which suggest environments closest to<br />

tropical African, non-seasonal, wet, evergreen<br />

forest faunas today, whereas the<br />

slightly younger (17·5–17·9 Ma) Rusinga<br />

<strong>and</strong> Mfangano Isl<strong>and</strong> sites were most similar<br />

to dry seasonal forests <strong>and</strong> also had more<br />

open conditions. This evidence may turn<br />

out to be important in considering information<br />

about whether the diets <strong>of</strong> these<br />

different species <strong>of</strong> Proconsul were similar<br />

<strong>and</strong> in interpreting the effects <strong>of</strong> seasonality<br />

on developing tooth tissues among different<br />

species <strong>of</strong> Proconsul (Macho et al., 1996).<br />

The paleosols also provide useful environmental<br />

information. Retallack et al. (1995)<br />

have associated Proconsul from Rusinga<br />

Isl<strong>and</strong> with soils interpreted as having supported<br />

riparian woodl<strong>and</strong> early in the ecological<br />

succession <strong>of</strong> streamsides. These<br />

workers found no evidence <strong>of</strong> soils that<br />

would indicate extensive dry grassl<strong>and</strong>s or<br />

wet rain forest. Substantial paleobotanical<br />

remains are found on Rusinga <strong>and</strong><br />

Mfangano. The type <strong>of</strong> P. heseloni was<br />

deposited by a predator in a large hollow<br />

tree (Walker & Teaford, 1988). Fossilized<br />

fruits have been found in the same paleosol<br />

as the partial skeletons, some teeth <strong>of</strong> which<br />

form the biggest sample for this study, at the<br />

Kaswanga Primate Site (Walker et al., 1985)<br />

<strong>and</strong> again these paleobotanical finds point to<br />

potential differences in seasonality <strong>and</strong> diet<br />

in Proconsul from Rusinga Isl<strong>and</strong>.<br />

On the basis <strong>of</strong> histological sections <strong>of</strong><br />

nine molar teeth attributed to P. africanus,<br />

P. major <strong>and</strong> P. nyanzae, Gantt (1983,<br />

1986), has previously reported that linear<br />

measurements <strong>of</strong> enamel indicate thick<br />

enamel, relative to body size estimates.<br />

Gantt estimated enamel thickness in these

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