Impacts of climate change on species, populations and communities: palaeobiogeographical insights and frontiers

Understanding climate change and its potential impact on species, populations and communities is one of the most pressing questions of twenty-fi rst-century conservation planning. Palaeobiogeographers working on Cenozoic fossil records and other lines of evidence are producing important insights into the dynamic nature of climate and the equally dynamic response of species, populations and communities. Climatic variations ranging in length from multimillennia to decades run throughout the palaeo-records of the Quaternary and earlier Cenozoic and have been shown to have had impacts ranging from changes in the genetic structure and morphology of individual species, population sizes and distributions, community composition to large-scale bio-diversity gradients. The biogeographical impacts of climate change may be due directly to the effects of alterations in temperature and moisture on species, or they may arise due to changes in factors such as disturbance regimes. Much of the recent progress in the application of palaeobiogegraphy to issues of climate change and its impacts can be attributed to developments along a number of still advancing methodological frontiers. These include increasingly finely resolved chronological resolution, more refi ned atmosphere-biosphere modelling, new biological and chemical techniques in reconstructing past species distributions and past climates, the development of large and readily accessible geo-referenced databases of biogeographical and climatic information, and new approaches in fossil morphological analysis and new molecular DNA techniques.

Variability in thermal and UV-B energy fluxes through time and their influence on plant diversity and speciation

Throughout Earth's history there have been temporal and spatial variations in the amount of visible and ultraviolet radiation received by ecosystems. This paper examines if temporal changes in these forms of energy receipt could have influenced the tempo and mode of plant diversity and speciation, focusing in particular upon Cenozoic time-scales. Evidence for changing patterns of plant diversity and speciation apparent in various fossil records and molecular phylogenies are considered alongside calculated changes in thermal and solar ultraviolet energy (specifically UV-B) over the past 50 Myr. We suggest that changes in thermal energy influx (amount and variability) affected the tempo of evolution through its influence upon community dynamics (e.g. population size, diversity, turnover, extinctions). It was not only the amount of thermal energy but also variability in its flux that may have influenced these processes, and ultimately the rate of diversification. We suggest that variations in UV-B would have influenced the mode and tempo of speciation through changes to genome stability during intervals of elevated UV-B. We argue, therefore, that although variability in thermal energy and UV-B fluxes through time may lead to the same end-point (changing the rate of diversification), the processes responsible are very different and both need to be considered when linking evolutionary processes to energy flux.

Neotropical refugia

Patterns of endemism in the Neotropics have been explained by restriction of forest to ‘refugia’ in arid cold-stages of the Quaternary (Haffer J (1969)
Speciation in Amazonian forest birds. Science 165: 131–137). The palaeoecological record, however, shows no such forest contraction. We review
palaeoecological and phylogenetic data on the response of Neotropical taxa and communities to climatic changes of the Cenozoic. Solar insolation varies
over this period with latitude and geography, including shifts in opposite directions between high and low latitudes. In the Neotropics, distribution and
abundance patterns originate on a wide range of timescales through the Cenozoic, down to the currently dominant precession forcing (20 kyr). In contrast,
distributions and abundances at higher latitudes are controlled by obliquity forcing (40 kyr). The patterns observed by Haffer (1969) are likely derived
from pre-Quaternary radiations and are not inconsistent with palaeoecological findings of continuous forest cover in major areas of the Neotropics
during the Quaternary. The relative proportions of speciation processes have changed through time between predominantly sympatric to predominantly
allopatric depending on the prevailing characteristics of orbitally forced climatic changes. Behaviour of Neotropical organisms and ecosystems on long
timescales may be influenced much more by precessional forcing than by the obliquity forcing that controls high-latitude climate change and glaciations.

The Klondike goldfields and Pleistocene environments of Beringia

The Klondike goldfields of Yukon, Canada, contain a key record of Pleistocene Beringia, the region of Alaska, Siberia, and Yukon that remained largely unglaciated during the late Cenozoic. A concentration of mining exposures, with relict permafrost that is locally more than 700,000 years old, provides exceptional preservation of paleoenvironmental archives and a new perspective on the nature of paleoenvironments during the Pleistocene. A critical feature is the stratigraphic association of distal tephra beds with these paleoenvironmental archives, which facilitates their regional correlation and, in many cases, provides independent ages for the paleoenvironmental assemblages. Paleoenvironmental analyses of fossil arctic ground-squirrel middens and buried vegetation indicate the presence of cryoxerophilous ("steppe-tundra") vegetation growing on well-drained substrates with deep active layers (seasonal thaw depths) during cold intervals of the Pleistocene. Studies of full-glacial paleosols and cryostratigraphic relations of associated ground ice indicate the importance of active loess deposition and surface vegetation cover in maintaining the functionally distinct mammoth-steppe biome, which supported grazing mega-fauna populations, including mammoth, horse, and bison.