The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism

The concept of ‘Gondwana’, an ancient Southern Hemisphere supercontinent, is firmly established in geological and biogeographical models of Earth history. The term Gondwana (Gondwanaland of some authors) derives from the recognition by workers at the Indian Geological Survey in the mid- to late 19th century of a distinctive sedimentary sequence preserved in east central India. This succession, now known to range in age from Permian to Cretaceous, is lithologically and palaeontologically similar to coeval non-marine sedimentary successions developed in most of the Southern Hemisphere continents suggesting former continuity of these landmasses. Palaeomagnetic data and tectonic reconstructions suggest that the main assembly of Gondwana took place around the beginning of the Palaeozoic in near-equatorial latitudes and that the supercontinent as a whole shifted into high southern latitudes, allowing widespread glaciation by the end of the Carboniferous. From Carboniferous to Cretaceous times the southern continents had broadly similar floras but some species-level provincialism is apparent at all times. The break-up of Gondwana initiated during the Jurassic (at about 180 million years ago) and this process is continuing. The earliest rifting (crustal attenuation) within the supercontinent initiated in the west (between South America and Africa) and in general terms the rifting pattern propagated eastward with major phases of continental fragmentation in the Early Cretaceous and Late Cretaceous to Paleogene. Gondwanan floras show radical turnovers near the end of the Carboniferous, end of the Permian and the end of the Triassic that appear to be unrelated to isolation or fragmentation of the supercontinent. Throughout the late Palaeozoic and Mesozoic the high-latitude southern floras maintained a distinctly different composition to the palaeoequatorial and boreal regions even though they remained in physical connection with Laurasia for much of this time. Gondwanan floras of the Jurassic and Early Cretaceous (times immediately preceding and during break-up) were dominated by araucarian and podocarp conifers and a range of enigmatic seed-fern groups. Angiosperms became established in the region as early as the Aptian (before the final break-up events) and steadily diversified during the Cretaceous, apparently at the expense of many seed-fern groups. Hypotheses invoking vicariance or long distance dispersal to account for the biogeographic patterns evident in the floras of Southern Hemisphere continents all rely on a firm understanding of the timing and sequence of Gondwanan continental breakup. This paper aims to summarise the current understanding of the geochronological framework of Gondwanan breakup against which these biogeographic models may be tested. Most phytogeographic studies deal with the extant, angiosperm-dominated floras of these landmasses. This paper also presents an overview of pre-Cenozoic, gymnosperm-dominated, floristic provincialism in Gondwana. It documents the broad succession of pre-angiosperm floras, highlights the distinctive elements of the Early Cretaceous Gondwanan floras immediately preceding the appearance of angiosperms and suggests that latitudinal controls strongly influenced the composition of Gondwanan floras through time even in the absence of marine barriers between Gondwana and the northern continents.