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Terrestrial Palaeoecology and Global Change

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Chapter 7. Climate change<br />

201<br />

are evolutionary conservative organs, the more leaf characters are indiscriminately used<br />

as past climate indicators, the greater a potential there is for an error to occur as a result<br />

of such misleading similarities. This consideration casts some doubt on the alleged advantage<br />

of using many poorly understood characters instead of few better known ones.<br />

Reproductive characters. Some reproductive features, such as volume-regulating<br />

(harmomegathy) structures in pollen grains, are commonly interpreted as climatic adaptations.<br />

However, an importance of harmomegathy primarily depends on pollination mode,<br />

with air-borne pollen grains normally exposed to desiccation for a shorter time than those<br />

carried by insects. Generally, over the evolutionary history of reproductive structures,<br />

the climatic effects are indirect, related to reproductive strategies. Thus, wind pollination<br />

prevails in temperate climate because of relatively low biological diversity, high population<br />

densities <strong>and</strong> the seasonal leaf shedding (canopies in full leaf screen the pollen rain).<br />

Entomophily is more frequent in tropical climates for the opposite reasons.<br />

Fossil evidence of pollination ecology is an important, although as yet little explored,<br />

source of palaeoclimatological inference. For example, the giant male strobili of Cyc<strong>and</strong>ra,<br />

a Jurassic cycadophyte (Fig. 88), produced thick-walled sporangia with narrow<br />

apical beaks, from which pollen grains, several hundreds per sporangium, were released<br />

in small portions. In contrast to instantaneous dispersal of wind-born pollen, this peculiar<br />

mechanism of parsimonious exposure offered a persistent pollen source for diachronously<br />

developing ovulate cones implying an extended pollination period that in turn implies<br />

a weak seasonality or none at all, with pollinators available round the year (Krassilov,<br />

2000a).<br />

More than one pollen type in gut compressions of fossil insects is evidence of overlapping<br />

pollination periods in the forage species. Such mixed pollen loads are rather<br />

typical of the mid-Permian insects of Cisuralia (Krassilov & Rasnitsyn, 1997; Krassilov<br />

et al., 1999d). The pollen types of their gut assemblages are also abundant in contemporaneous<br />

dispersed assemblages indicating the dominant pollen producers. This type plant<br />

– insect interactions indicates seasonality, with synchronous pollination probably confined<br />

to the drier season. In contrast, all hitherto studied Early Cretaceous xyelids from<br />

Baisa, Transbaikalia, had single-species pollen loads, with pollen types produced by relatively<br />

rare plant species. Thus the feeding habits were highly selective <strong>and</strong> probably<br />

adapted to diachronous pollen sources, as in the case of Ceroxyella dolichocera that<br />

fed either on saccate pollen produced by proangiospermous Preflosella or on cryptosaccate<br />

pollen of an unknown relative of the latter (VIII.3.3). Since xyelids are early<br />

spring insects, the succession of flowering should have been rapid as in ephemeroids of<br />

temperate climates.<br />

Like fossil pollen, fossil disseminules are as yet a poorly explored source of palaeoclimatic<br />

inference. The seed/fruit size is a function of (1) growth rates, with larger disseminules<br />

facilitating a faster seedling growth; (2) seed dormancy period correlated with<br />

small endosperm <strong>and</strong> thick coats; (3) seral stage, with disseminules typically smaller in<br />

pioneer species <strong>and</strong> colonizers (Wilson, 1993), (4) dispersal mode, in particular, the body

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