Grasses, by their high productivity even under very low pCO2, their ability to survive repeated burning and to tolerate long dry seasons, have transformed the terrestrial biomes in the Neogene and Quaternary. The expansion of grasslands at the cost of biodiverse forest biomes in Madagascar is often postulated as a consequence of the Holocene settlement of the island by humans. However, we show that the Malagasy grass flora has many indications of being ancient with a long local evolutionary history, much predating the Holocene arrival of humans. First, the level of endemism in the Madagascar grass flora is well above the global average for large islands. Second, a survey of many of the more diverse areas indicates that there is a very high spatial and ecological turnover in the grass flora, indicating a high degree of niche specialization. We also find some evidence that there are both recently disturbed and natural stable grasslands: phylogenetic community assembly indicates that recently severely disturbed grasslands are phylogenetically clustered, whereas more undisturbed grasslands tend to be phylogenetically more evenly distributed. From this evidence, it is likely that grass communities existed in Madagascar long before human arrival and so were determined by climate, natural grazing and other natural factors. Humans introduced zebu cattle farming and increased fire frequency, and may have triggered an expansion of the grasslands. Grasses probably played the same role in the modification of the Malagasy environments as elsewhere in the tropics.
Recent developments in grass taxonomy give a new insight into their classification, and point to a phylogenetic sequence which maps differences in their internal metabolism. Corroboratory fossil evidence is unfortunately exceedingly meager, but it can be supplemented by examining the implications of present-day distributions. The subfamilies are distributed in worldwide climatic belts, but two-thirds of the genera are confined to single continents. Obviously, the genera are poor travellers, so how did the grasses become so widespread? Much depends on the probability of transoceanic transport. The evidence is inconclusive, but it seems likely that the tropical subfamilies spread during the first half of the Tertiary when the maximum water gap was 1200 km. Species distributions are likewise influenced by climatic differentiation and continental isolation. But they sometimes reveal the intervention of other factors, particularly the disruptive effect of climatic change in the Pleistocene. Data from the Afro-montane flora do not support the proposition that species from adjacent, but contrasting, ecological environments are distributed independently. Nor does a mapping of endemics encourage the concept of discrete centers.
At altitudes below 1300 m most species of grasses in tropical Africa are ones that use 4-carbon photosynthesis. Above 4000 m only species of the subfamily Pooideae using 3-carbon photosynthesis are found. At intermediate altitudes the percent of 3-carbon genera and species in the grass flora is a very regular function of altitude. The correlate of altitude that controls the distribution of grasses appears to be temperature. Fossil grass cuticles are identifiable to genus and should provide a useful paleothermometer. If no other errors were involved, the regular altitudinal distribution of genera in the flora of tropical East Africa would permit paleotemperature estimates with 95% confidence limits of ± 1.2°C.
Aim For most higher‐order taxa, species diversity peaks sharply in the moist tropics and declines rapidly at higher latitudes. However, the mechanisms driving this latitudinal gradient are numerous, remain uncertain and are even undocumented in some important major clades such as the grasses. Grasses are a cosmopolitan, important plant family with more than 11,000 species world‐wide. Our aims were: to investigate the latitudinal distributions of species richness for different grass lineages, and the grass family as a whole; and to test hypotheses proposed in general for the latitudinal diversity gradient or specifically as determinants of grass species richness patterns at the global scale. Location Global. Methods We used the most comprehensive global database of grass distributions currently available to calculate species richness for 340 political regions of the world. Using generalized additive models we used these data to model the latitudinal gradients of species richness for different grass lineages and for the grass family as a whole. We constructed multiple regression models to include climatic, productivity, topographic, habitat and geographic variables. Results An unusual, shallow latitudinal diversity gradient arises because different grass lineages exhibit contrasting latitudinal patterns. This reflects differential specialization of grass lineages to arid and cool environments, the legacy of historical effects, most notably the Gondwanan origin of the grasses, and a strong association between grasses and topographically heterogeneous, mountainous regions. Main conclusions The grasses are one of the relatively few higher‐order lineages that exhibits an atypical latitudinal gradient; this has arisen because of climatic specialization of particular grass lineages to cold and arid environments. Key additional roles have been inferred for historical biogeography and topographical heterogeneity in determining global patterns of grass species richness. These mechanisms have generally been under‐appreciated and are probably important for many other lineages.
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