Reproductive strategy and the life cycle in graptoloid colonies

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Abstract
Graptoloid colonies were clones composed in all probability of hermaphroditic zooids. Their breeding system approaching that of amphicarpic plants, namely distant outcrossing combined with selfing, was exceptionally flexible. Moreover, the balanced coexistence of these extreme means of reproduction may be visualized as an evolutionarily stable strategy (ESS). The evolutionary consequences of such a breeding system might have accounted for the overall high rates of graptoloid evolution as well as for the rapid transformations in large populations. Sexual reproduction of graptoloid colonies was complemented by a multiplication through occasional fragmentation of colonies and subsequent regeneration from the fragments. Fragmentation of colonies played an important role in the survival of graptoloid colonies during catastrophic events such as hurricanes and later in recruitment. Fragmentation followed by regeneration, and only later by sexual reproduction of regenerated fragments, constituted the so-called great cycle (GC), whereas the regular course of events initiated by sexual reproduction in an undisturbed (complete) colony resulted in normal colony formation (astogeny) and is termed the small cycle (SC). Thus the adaptive significance of the sexual process in the life cycle is in the restoration of the 'perfect' pattern of the complete colony, which offered the best hydrodynamic properties and highest fitness.