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Comparative dental development and microstructure of ... - UCL

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

from the estimated mean age <strong>of</strong> M1 emergence<br />

based on data for brain size <strong>and</strong> M1<br />

emergence in 23 species <strong>of</strong> primates (Kelley,<br />

1997). In P. heseloni, M1 emergence was<br />

estimated at 20·6 months with an approximate<br />

range for the mean (based on the<br />

confidence interval for brain size rather than<br />

the error <strong>of</strong> the estimate) <strong>of</strong> between 19·6<br />

<strong>and</strong> 21·6 months. Kelley (1997) concluded<br />

that this age for M1 emergence falls at the<br />

upper end <strong>of</strong> the range <strong>of</strong> means for all<br />

extant nonhominoid catarrhines, many <strong>of</strong><br />

which are considerably larger on average<br />

than P. heseloni. On this evidence Kelley<br />

(1997) has cautiously suggested these<br />

results indicate a more prolonged life history<br />

for P. heseloni. But a considerable grade shift<br />

may occur between Old World monkeys <strong>and</strong><br />

hominoids in many life history traits <strong>and</strong> in<br />

<strong>dental</strong> <strong>development</strong> <strong>and</strong> predictions based<br />

only on hominoid brain sizes, for example,<br />

might result in estimates <strong>of</strong> M1 emergence<br />

in P. heseloni in excess <strong>of</strong> those cited here.<br />

Estimates for M 1 crown formation time in<br />

P. heseloni in this study are around 14<br />

months after birth (approximately 30 days <strong>of</strong><br />

M 1 formation occurred before birth). If root<br />

extension occurred at the same rate as in M 2<br />

<strong>and</strong> P 4 at an average 6·4 μm per day (<strong>and</strong><br />

poor data from one M 1 section suggests this<br />

is likely) then M1 would have just less than<br />

1·25 mm <strong>of</strong> root formed at 20·6 months but<br />

close to 1·5 mm formed at 21·6 months.<br />

These predictions are speculative but not<br />

incompatible with the predictions <strong>of</strong> Kelley<br />

(1997) based on 23 species <strong>of</strong> primates.<br />

Future studies on <strong>dental</strong> <strong>development</strong> in P.<br />

nyanzae <strong>and</strong> P. major might result in good<br />

estimates for the age <strong>of</strong> emergence <strong>of</strong> M1 in<br />

these larger bodied proconsulids <strong>and</strong> so provide<br />

a better idea about the affects <strong>of</strong> body<br />

size <strong>and</strong> tooth size on <strong>dental</strong> <strong>development</strong><br />

within Proconsul. This would make it easier<br />

to judge whether there is evidence that<br />

<strong>dental</strong> <strong>development</strong> <strong>and</strong> the overall maturational<br />

pr<strong>of</strong>ile was prolonged in Proconsul.<br />

Histological studies on other Miocene primates<br />

such as Victoriapithecus, Sivapithecus<br />

<strong>and</strong> Lufengpithecus will also place the results<br />

<strong>of</strong> this study into a better phylogenetic<br />

perspective. It remains likely, however, that<br />

postcranial, masticatory <strong>and</strong> life history<br />

traits evolved in a mosaic fashion (Rae,<br />

1997). Of these traits, those that probably<br />

resulted from reduced adult mortality rates<br />

(which include a prolonged <strong>development</strong>al<br />

period <strong>and</strong> a bigger brain) are likely to have<br />

been the last to appear.<br />

Conclusions<br />

This is the first histological study <strong>of</strong><br />

Proconsul teeth from Rusinga Isl<strong>and</strong>, Kenya.<br />

A chronology <strong>of</strong> the sequence <strong>of</strong> tooth<br />

<strong>development</strong> in P. heseloni indicates M3 root<br />

formation was complete between 6 <strong>and</strong><br />

7 years in this siamang-sized Miocene<br />

primate. Crown formation times in an M 1<br />

<strong>and</strong> M 2 attributed to the larger female<br />

chimpanzee-sized P. nyanzae were between<br />

30% <strong>and</strong> 40% less than the average values<br />

for seven common chimpanzees studied in<br />

the same way. The results reported here<br />

suggest that both species <strong>of</strong> Proconsul from<br />

Rusinga Isl<strong>and</strong> had thicker enamel than previously<br />

described for P. africanus <strong>and</strong> P.<br />

major from other older sites in western<br />

Kenya <strong>and</strong> Ug<strong>and</strong>a. Reports on the palaeoecology<br />

<strong>of</strong> Miocene sites on Rusinga<br />

together with future research on the evidence<br />

for seasonality from accentuated<br />

markings in teeth <strong>and</strong> from tooth microwear<br />

studies may allow us to place studies <strong>of</strong><br />

enamel thickness in Proconsul into a more<br />

secure dietary <strong>and</strong> functional context.<br />

Certain microstructural features in enamel<br />

<strong>and</strong> dentine appear, so far, to be unique to<br />

Proconsul. P. nyanzae appears more derived<br />

with respect to P. heseloni in the degree to<br />

which these unique features are expressed.<br />

Rates <strong>of</strong> enamel formation close to the EDJ<br />

are higher than in other primates studied so<br />

far. The ratio <strong>of</strong> dentine to enamel formed<br />

close to the EDJ in the lateral aspects <strong>of</strong> the

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