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Evolution__3rd_Edition

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Incidence of repair<br />

0.6<br />

0.5<br />

0.4<br />

0.3<br />

0.2<br />

0.1<br />

0.0<br />

50<br />

40<br />

30<br />

20<br />

10<br />

0<br />

Late<br />

Carboniferous<br />

5–9 mm<br />

10–19 mm<br />

20–29 mm<br />

Late Late<br />

Triassic Cretaceous<br />

Time interval<br />

. . . in fossil–predator prey relations<br />

...<br />

Percentage of occurrence of<br />

uncemented gastropods<br />

Late<br />

Miocene<br />

Recent<br />

CHAPTER 22 / Coevolution 635<br />

Figure 22.12<br />

Incidence of repair at five successive time periods, for shells<br />

divided into three size classes. Note: (i) the incidence of repair is<br />

higher in more recent times; and (ii) larger shells show a relatively<br />

high incidence of repair compared with small shells in more<br />

recent times, as compared with earlier times. Large shells tend to<br />

be more resistant to breakage than small ones. One interpretation<br />

of the second trend is that more recent predators have become<br />

stronger, and therefore able to injure large-shelled animals.<br />

(0.4 in ≈ 10 mm.) Redrawn, by permission of the publisher, from<br />

Vermeij (1987).<br />

limited evidence. Those broad patterns, as we shall see, have a wide scatter in the data.<br />

Dietl et al.’s point is worth keeping in mind, because problems in the data are likely to<br />

be one cause of the scatter.<br />

Let us look at some of Vermeij’s evidence. The frequency of shell repair is one indicator<br />

of predator–prey interactions in fossils. When a mollusk is non-lethally attacked,<br />

it repairs the damage to its shell and the repair pattern can be observed in the shell.<br />

Proportions of shells showing signs of repair have been measured in several fossil<br />

faunas, and the trend appears to be toward increasing amounts of repair over time<br />

(Figure 22.12). This Vermeij interprets as meaning that the prey have been suffering<br />

higher frequencies of predatory attacks over evolutionary time. (Logically it could also<br />

mean a though this is perhaps unlikely a that the predators have de-escalated from<br />

forms that destroyed their prey to forms that sometimes merely injured them!)<br />

The escalation of molluskan prey defenses is also suggested by a trend in the proportion<br />

of different types of gastropod shells through time. The proportion of loosely<br />

attached forms, which are relatively poorly defended, has decreased through time relative<br />

to better defended types, such as burrowers and attached forms (Figure 22.13).<br />

Vermeij also found limited evidence that the better defended burrowers increased from<br />

being about 5–10% of genera in the late Carboniferous–late Triassic to about 37%<br />

in the late Cretaceous and 62–75% in modern formations. Internal thickening or narrowing<br />

of the aperture is another form of escalated defense and these types, too, have<br />

(2) (3) (3)<br />

(2)<br />

Ord. Sil. Dev. E. L. Per. Tri. Jur. Cret. Neog.<br />

Carb. Carb.<br />

Time interval<br />

Figure 22.13<br />

The incidence of sessile or sedentary uncemented gastropods<br />

through time. Note that the proportion decreases. Each point<br />

is for one fossil assemblage, except where marked. For each<br />

assemblage, Vermeij divided the gastropods into different types<br />

(burrowers, sessile, attached forms, etc.), adding up to 100%. This<br />

graph gives the proportion of gastropods that lie unattached on<br />

the bottom surface. The Neogene (Neog.) includes the Miocene<br />

and Pliocene (Figure 18.1, p. 526). Redrawn, by permission of the<br />

publisher, from Vermeij (1987).

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