The antiquity of Madagascar’s grasslands and the rise of C4 grassy biomes

Aim Grasslands and savannas, which make up > 75% of Madagascar’s land area, have long been viewed as anthropogenically derived after people settled on the island c. 2 ka. We investigated this hypothesis and an alternative – that the grasslands are an insular example of the post‐Miocene spread of C₄ grassy biomes world‐wide. Location Madagascar, southern Africa, East Africa. Methods We compared the number of C₄ grass genera in Madagascar with that in southern and south‐central African floras. If the grasslands are recent we would expect to find fewer species and genera in Madagascar relative to Africa and for these species and genera to have very wide distribution ranges in Madagascar. Secondly, we searched Madagascan floras for the presence of endemic plant species or genera restricted to grasslands. We also searched for evidence of a grassland specialist fauna with species endemic to Madagascar. Plant and animal species endemic to C₄ grassy biomes would not be expected if these are of recent origin. Results Madagascar has c. 88 C₄ grass genera, including six endemic genera. Excluding African genera with only one or two species, Madagascar has 86.6% of southern Africa’s and 89.4% of south‐central Africa’s grass genera. C₄ grass species make up c. 4% of the flora of both Madagascar and southern Africa and species : genus ratios are similar (4.3 and 5.1, respectively). Turnover of grasses along geographical gradients follows similar patterns to those in South Africa, with Andropogoneae dominating in mesic biomes and Chlorideae in semi‐arid grassy biomes. At least 16 monocot genera have grassland members, many of which are endemic to Madagascar. Woody species in frequently burnt savannas include both Madagascan endemics and African species. A different woody flora, mostly endemic, occurs in less frequently burnt grasslands in the central highlands, filling a similar successional niche to montane C₄ grasslands in Africa. Diverse vertebrate and invertebrate lineages have grassland specialists, including many endemic to Madagascar (e.g. termites, ants, lizards, snakes, birds and mammals). Grassland use of the extinct fauna is poorly known but carbon isotope analysis indicates that a hippo, two giant tortoises and one extinct lemur ate C₄ or CAM (crassulacean acid metabolism) plants. Main conclusions The diversity of C₄ grass lineages in Madagascar relative to that in Africa, and the presence of plant and animal species endemic to Madagascan grassy biomes, does not fit the view that these grasslands are anthropogenically derived. We suggest that grasslands invaded Madagascar after the late Miocene, part of the world‐wide expansion of C₄ grassy biomes. Madagascar provides an interesting test case for biogeographical analysis of how these novel biomes assembled, and the sources of the flora and fauna that now occupy them. A necessary part of such an analysis would be to establish the pre‐settlement extent of the C₄ grassy biomes. Carbon isotope analysis of soil organic matter would be a feasible method for doing this.     

Phylogenetic signals in the realized climate niches of Chinese grasses (Poaceae)

Explaining relationships between species richness and biogeographical patterns over a broad geographic scale is a central issue of biogeography and macroecology. We document the realized climate niches for grasses in China’s nature reserves and discuss its formation mechanism using grass richness data combined with climatic, physiological, and phylogenetic data. Our results suggest that climate niche structure of grasses is phylogenetically conservative for BEP (Bambusoideae, Ehrhartoideae, and Pooideae) and PACMAD (Panicoideae, Arundinoideae, Chloridoideae, Micrairoideae, Aristidoideae, and Danthonioideae) clades along temperature gradients and for Chloridoideae and Panicoideae along precipitation gradients. At the national scale, the divergence patterns of climate niches between two major clades are more distinguishable than between C₃ and C₄ grasses. High rates of climate niche evolution are found in C₄ clades in the subtropical forest region. There appears to be a strong association between elevation gradients and grass diversity: the specific environmental conditions (e.g. energy) and the rapid shifts of climate conditions drive high grass diversification. Evolutionary conservatism of climate niches may be influenced by the specific adaptive ability to changing environmental conditions within NAD-ME/NADP-ME clades. Our results indicate that adaptations to major climate changes may be accomplished by C₄ grass nodes of high climate niche evolutionary rates in China’s nature reserves.     

Phylogenetic niche conservatism in C4 grasses

Photosynthetic pathway is used widely to discriminate plant functional types in studies of global change. However, independent evolutionary lineages of C₄ grasses with different variants of C₄ photosynthesis show different biogeographical relationships with mean annual precipitation, suggesting phylogenetic niche conservatism (PNC). To investigate how phylogeny and photosynthetic type differentiate C₄ grasses, we compiled a dataset of morphological and habitat information of 185 genera belonging to two monophyletic subfamilies, Chloridoideae and Panicoideae, which together account for 90 % of the world’s C₄ grass species. We evaluated evolutionary variance and covariance of morphological and habitat traits. Strong phylogenetic signals were found in both morphological and habitat traits, arising mainly from the divergence of the two subfamilies. Genera in Chloridoideae had significantly smaller culm heights, leaf widths, 1,000-seed weights and stomata; they also appeared more in dry, open or saline habitats than those of Panicoideae. Controlling for phylogenetic structure showed significant covariation among morphological traits, supporting the hypothesis of phylogenetically independent scaling effects. However, associations between morphological and habitat traits showed limited phylogenetic covariance. Subfamily was a better explanation than photosynthetic type for the variance in most morphological traits. Morphology, habitat water availability, shading, and productivity are therefore all involved in the PNC of C₄ grass lineages. This study emphasized the importance of phylogenetic history in the ecology and biogeography of C₄ grasses, suggesting that divergent lineages need to be considered to fully understand the impacts of global change on plant distributions.     

The distribution of C3 and C4 grasses and carbon isotope discrimination along an altitudinal and moisture gradient in Kenya

Summary More than 500 species of the Poaceae are found in Kenya, East Africa. Eighteen of twenty-seven tribes are exclusively (except the Paniceae and Danthonieae) of the C₃ photosynthetic type. A floristic analysis of low altitude grasslands suggests that nearly all species at these low altitudes are of the C₄ photosynthetic type. At high altitudes, however, nearly all grasses are of the C₃ photosynthetic type. Open grassland vegetation was sampled along a transect from arid low altitude sites to the top of Mt. Kenya in an attempt to document the general distributions of the photosynthetic types.The major tribes illustrated three general patterns of distribution. The C₄ tribes Chlorideae, Eragrosteae, Sporoboleae, and Aristideae were abundant at low altitudes (or low indices of available soil moisture). The Paniceae and Andropogoneae were also exclusively C₄ but were more common at intermediate altitudes. The C₃ tribes Aveneae, Festuceae, and Agrostideae were found only at high altitudes. In these open grasslands there were no C₃ species below 2,000 m and no C₄ species above 3,000 m. The variation in ₁₃C of the live grass vegetation with altitude confirms these distributional patterns and suggests a sharp transition zone between these two photosynthetic types. The photosynthetic type accounts for broad distributions within the Poaceae but these distributions are further modified by characteristics which may be inherent in the tribal groups. Ecological and paleoecological significance of these patterns of distribution are discussed.     

When is a ‘forest’ a savanna,and why does it matter?

Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high‐rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C₄ grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C₄ grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life‐history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire‐resistant tree species in a C₄ grass‐dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.     

Preference for C4 shade grasses increases hatchling performance in the butterfly,Bicyclus safitza

The Miocene radiation of C₄ grasses under high‐temperature and low ambient CO₂ levels occurred alongside the transformation of a largely forested landscape into savanna. This inevitably changed the host plant regime of herbivores, and the simultaneous diversification of many consumer lineages, including Bicyclus butterflies in Africa, suggests that the radiations of grasses and grazers may be evolutionary linked. We examined mechanisms for this plant–herbivore interaction with the grass‐feeding Bicyclus safitza in South Africa. In a controlled environment, we tested oviposition preference and hatchling performance on local grasses with C₃ or C₄ photosynthetic pathways that grow either in open or shaded habitats. We predicted preference for C₃ plants due to a hypothesized lower processing cost and higher palatability to herbivores. In contrast, we found that females preferred C₄ shade grasses rather than either C₄ grasses from open habitats or C₃ grasses. The oviposition preference broadly followed hatchling performance, although hatchling survival was equally good on C₄ or C₃ shade grasses. This finding was explained by leaf toughness; shade grasses were softer than grasses from open habitats. Field monitoring revealed a preference of adults for shaded habitats, and stable isotope analysis of field‐sampled individuals confirmed their preference for C₄ grasses as host plants. Our findings suggest that plant–herbivore interactions can influence the direction of selection in a grass‐feeding butterfly. Based on this work, we postulate future research to test whether these interactions more generally contribute to radiations in herbivorous insects via expansions into new, unexploited ecological niches.     

Speciation and radiations track climate transitions since the Miocene Climatic Optimum: a case study of southern African chameleons

Aim The high amount of species diversity concentrated in southern Africa has been attributed to palaeoclimatic factors, and the timing of radiations in some taxa corresponds to global palaeoclimatic trends. Using dwarf chameleons (Bradypodion: Chamaeleonidae) as a model system, we explored the relationship between palaeoclimatic fluctuations and cladogenesis with respect to both temporal and spatial patterns in an effort to understand the process of speciation in southern Africa. Location South Africa, with particular emphasis on the Cape Floristic Region and the Maputaland–Pondoland–Albany hotspot. Methods Mitochondrial sequence data (ND2 and 16S) were used to estimate the timing of major radiations and to examine the number of lineages through time. A dated phylogeny was constructed using Bayesian phylogenetic reconstruction, and a Bayesian relaxed molecular clock was used to estimate divergence times. Spatial data and lineage‐through‐time plots were used to identify geographic regions that underwent diversification in connection with major climatic events. Both parsimony and likelihood optimizations of habitat type on the phylogeny were used to determine whether major habitat shifts have occurred. On a coarse scale (half‐degree grid cells), phylogenetic diversity (sum of the branch lengths linking terminals) was compared with species richness (absolute number of species) to identify areas of conservation importance. Results The complete species phylogeny of dwarf chameleons shows that the timing and mode of diversification exhibit spatio‐temporal patterns that link to phases in the evolution of southern Africa’s climate over the last 14 Myr. Optimizations of habitat on the phylogenetic tree show a progression from closed to open habitats since the Mid‐Miocene, corresponding to the shift from C₃ to C₄ environments, and later with the development of south‐western Africa’s winter‐rainfall regime. These shifts are not simultaneous across the region, with different geographic centres of diversity generated during different time periods. Main conclusions Regions that are prominent centres of chameleon diversification are encompassed by the current biodiversity hotspots as shown by chameleon species richness and phylogenetic diversity. Diversity within the Cape Floristic Region appears to be the result of a Late Pliocene radiation, whereas the diversity encompassed within the Maputaland–Pondoland–Albany hotspot is an aggregate of asynchronous radiation events, probably influenced by lineage losses. Overall, dwarf chameleons have experienced a shift in habitat types, with recent radiations occupying open habitats, and older lineages persisting in relictual forested habitats, corresponding to the continental shift of vegetation types since the Miocene Climatic Optimum.     

Photosynthetic pathway variation among C4 grasses along a precipitation gradient in Argentina

Aim Based on the biochemical and physiological attributes of C₄ grasses, and on the close association between decarboxylation pathways and the taxa in which they evolved, the hypotheses tested were: (1) that C₄ grasses would become progressively more abundant as precipitation decreased, with grasses of the NADP‐me subtype more abundant in wetter sites and those of the NAD‐me subtype more common in arid regions; and (2) that the distribution of grass subfamilies would also be correlated with annual precipitation. Location The study was conducted along a precipitation gradient in central Argentina, from the eastern Pampas (>1000 mm year^?1) to the western deserts and semi‐deserts near the Andes (<100 mm year^?1). Methods Percentage of species and relative cover of C₃ and C₄ grasses (including C₄ subtypes) in local floras from 15 lowland sites of central Argentina were obtained from our own unpublished data and from recently published floristic surveys. Pearson correlation coefficients were obtained between grass distribution parameters and the available climatic data. Results The percentage of C₄ grasses increased towards the arid extreme and showed a strong negative correlation with annual rainfall (r = ?0.74, P < 0.01). Within the C₄ subtypes, the NADP‐me species showed a higher proportional representation at the wetter extreme, whereas the representation of NAD‐me species increased towards the more arid extreme. The relationship of PEP‐ck species with climatic parameters in central Argentina was less evident. The distributions of the Panicoideae and Chloridoideae subfamilies along the precipitation gradient were diametrically opposed, with the Panicoideae positively (r = 0.86, P < 0.001) and the Chloridoideae negatively (r = ?0.87, P < 0.001) correlated with annual precipitation. Main conclusions Our data are consistent with the broad observation that C₄ grasses tend to dominate in areas where the wet season falls in the warmer summer months. In agreement with previously reported results for Africa, Asia, Australia and North America, we describe here for the first time a significant relationship between annual precipitation and the prevalence of the NADP‐me and NAD‐me photosynthetic pathways along climatic gradients for the Neotropics. We also report for the first time that correlations between C₄ species and annual rainfall are stronger when the relative cover of grass species is considered. The association of grass subfamilies Panicoideae and Chloridoideae with rainfall is as strong as that recorded for the NADP‐me and NAD‐me variants, respectively, suggesting that characteristics other than decarboxylation type may be responsible for the geographic patterns described in this study.     

Cascading biodiversity and functional consequences of a global change–induced biome switch

Aim At a regional scale, across southern Africa, woody thickening of savannas is becoming increasingly widespread. Using coupled vegetation and faunal responses (ants), we explore whether major changes in woody cover in savannas represent an increase in the density of savanna trees (C₄ grass layer remains intact) or a ‘regime shift’ in system state from savanna to thicket (=dry forest) where broad‐leaved, forest‐associated trees shade out C₄ grasses. Location Hluhluwe Game Reserve, South Africa. Methods We sampled paired open (low woody cover) and closed (high cover that have undergone an increase in tree density) sites. Vegetation was sampled using belt transects, and a combination of pitfall trapping and Winkler sampling was used for ants. Results Closed habitats did not simply contain a higher density of woody savanna species, but differed significantly in structure, functional composition (high prevalence of broad‐leaved trees, discontinuous C₄ grasses) and system properties (e.g. low flammability). Ant assemblage composition reflected this difference in habitat. The trophic structure of ant assemblages in the two habitats revealed a functional shift with much higher abundances of predatory species in the closed habitat. Main conclusions The predominance of species with forest‐associated traits and concomitant reduction of C₄ grasses in closed sites indicate that vegetation has undergone a shift in fundamental system state (to thicket), rather than simply savanna thickening. This biome shift has cascading functional consequences and implications for biodiversity conservation. The potential loss of many specialist savanna plant species is especially concerning, given the spatial extent and speed of this vegetation switch. Although it is not clear how easily the habitat switch can be reversed and how stable the thicket habitats are, it is likely in the not‐too‐distant future that conservation managers will be forced to make decisions on whether to actively maintain savannas.     

Dominant Grasses Suppress Local Diversity in Restored Tallgrass Prairie

Warm‐season (C₄) grasses commonly dominate tallgrass prairie restorations, often at the expense of subordinate grasses and forbs that contribute most to diversity in this ecosystem. To assess whether the cover and abundance of dominant grass species constrain plant diversity, we removed 0, 50, or 100% of tillers of two dominant species (Andropogon gerardii or Panicum virgatum) in a 7‐year‐old prairie restoration. Removing 100% of the most abundant species, A. gerardii, significantly increased light availability, forb productivity, forb cover, species richness, species evenness, and species diversity. Removal of a less abundant but very common species, P. virgatum, did not significantly affect resource availability or the local plant community. We observed no effect of removal treatments on critical belowground resources, including inorganic soil N or soil moisture. Species richness was inversely correlated with total grass productivity and percent grass cover and positively correlated with light availability at the soil surface. These relationships suggest that differential species richness among removal treatments resulted from treatment induced differences in aboveground resources rather than the belowground resources. Selective removal of the dominant species A. gerardii provided an opportunity for seeded forb species to become established leading to an increase in species richness and diversity. Therefore, management practices that target reductions in cover or biomass of the dominant species may enhance diversity in established and grass‐dominated mesic grassland restorations.     

Macro-Climatic Distribution Limits Show Both Niche Expansion and Niche Specialization among C4 Panicoids

Grasses are ancestrally tropical understory species whose current dominance in warm open habitats is linked to the evolution of C₄ photosynthesis. C₄ grasses maintain high rates of photosynthesis in warm and water stressed environments, and the syndrome is considered to induce niche shifts into these habitats while adaptation to cold ones may be compromised. Global biogeographic analyses of C₄ grasses have, however, concentrated on diversity patterns, while paying little attention to distributional limits. Using phylogenetic contrast analyses, we compared macro-climatic distribution limits among ~1300 grasses from the subfamily Panicoideae, which includes 4/5 of the known photosynthetic transitions in grasses. We explored whether evolution of C₄ photosynthesis correlates with niche expansions, niche changes, or stasis at subfamily level and within the two tribes Paniceae and Paspaleae. We compared the climatic extremes of growing season temperatures, aridity, and mean temperatures of the coldest months. We found support for all the known biogeographic distribution patterns of C₄ species, these patterns were, however, formed both by niche expansion and niche changes. The only ubiquitous response to a change in the photosynthetic pathway within Panicoideae was a niche expansion of the C₄ species into regions with higher growing season temperatures, but without a withdrawal from the inherited climate niche. Other patterns varied among the tribes, as macro-climatic niche evolution in the American tribe Paspaleae differed from the pattern supported in the globally distributed tribe Paniceae and at family level.     

Distribution of South African C3 and C4 species of Cyperaceae in relation to climate and phylogeny

Abstract In this study the contribution of climatic factors and phylogenetic relationships affecting the geographical distribution of C₃ and C₄ genera of the Cyperaceae in South Africa was investigated. The δ¹³C values of herbarium specimens of 68 southern African species from 22 genera and eight tribes were used to assign the species to either the C₃ or C₄ photosynthetic pathway. Geographical distribution data for the Cyperaceae were used to investigate relationships between climatic factors and the number of species and proportional abundance of C₄ species per region. The number of Cyperaceae species per 2° × 2° square across South Africa varied from less than five in the north‐western regions to more than 15 in the south‐western and north‐eastern regions of South Africa where rainfall exceeds 800 mm y^‐1. Of the 68 species investigated, 28 had C₄ photosynthesis and these were scattered among nine genera of four tribes (Cypereae, Scirpeae, Abildgaardieae and Rhyncosporeae). The proportional abundance of C₄ species ranged from 14% in the winter rainfall regions of the south‐west of South Africa to 67% in the summer rainfall areas of the north‐east. The geographical distribution of species was related to their phylogenetic position such that the distributions of C₃ and C₄ species in Cypereae, Scirpeae and Schoeneae was quite distinct. Linear regression analysis showed that the transition temperatures (equal C₃ and C₄ species numbers) for the Cyperaceae were different to those obtained for the Poaceae from the same region. No strong relationships were found between the proportional abundance of C₄ species and other climate factors such as altitude and rainfall. Our analysis of the current geographical distribution of C₄ Cyperaceae in southern Africa in a phylogenetic context suggests that the ecological advantages conferred by the C₄ pathway differ amongst the different plant groups.     

Biogeography of the grasses (Poaceae): a phylogenetic approach to reveal evolutionary history in geographical space and geological time

The grasses (Poaceae) are the fifth most diverse family of angiosperms, including 800 genera and more than 10 000 species. Few phylogenetic studies have tried to investigate palaeo‐biogeographical and palaeo‐ecological scenarios that may have led to present‐day distribution and diversity of grasses at the family level. We produced a dated phylogenetic tree based on combined plastid DNA sequences and a comprehensive sample of Poaceae. Furthermore, we produced an additional tree using a supermatrix of morphological and molecular data that included all 800 grass genera so that ancestral biogeography and ecological habitats could be inferred. We used a likelihood‐based method, which allows the estimation of ancestral polymorphism in both biogeographical and ecological analyses for large data sets. The origin of Poaceae was retrieved as African and shade adapted. The crown node of the BEP + PACCMAD clade was dated at 57 Mya, in the early Eocene. Grasses dispersed to all continents by approximately 60 million years after their Gondwanan origin in the late Cretaceous. PACCMAD taxa adapted to open habitats as early as the late Eocene, a date consistent with recent phytolith fossil data for North America. C₄ photosynthesis first originated in Africa, at least for Chloridoideae in the Eocene at c. 30 Mya. The BEP clade members adapted to open habitats later than PACCMAD members; this was inferred to occur in Eurasia in the Oligocene. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 543–557.     

Variations in nitrogen use efficiency reflect the biochemical subtype while variations in water use efficiency reflect the evolutionary lineage of C4 grasses at inter‐glacial CO2

C₄ photosynthesis evolved multiple times in diverse lineages. Most physiological studies comparing C₄ plants were not conducted at the low atmospheric CO₂ prevailing during their evolution. Here, 24 C₄ grasses belonging to three biochemical subtypes [nicotinamide adenine dinucleotide malic enzyme (NAD‐ME), phosphoenolpyruvate carboxykinase (PCK) and nicotinamide adenine dinucleotide phosphate malic enzyme (NADP‐ME)] and six major evolutionary lineages were grown under ambient (400 μL L^?1) and inter‐glacial (280 μL L^?1) CO₂. We hypothesized that nitrogen‐related and water‐related physiological traits are associated with subtypes and lineages, respectively. Photosynthetic rate and stomatal conductance were constrained by the shared lineage, while variation in leaf mass per area (LMA), leaf N per area, plant dry mass and plant water use efficiency were influenced by the subtype. Subtype and lineage were equally important for explaining variations in photosynthetic nitrogen use efficiency (PNUE) and photosynthetic water use efficiency (PWUE). CO₂ treatment impacted most parameters. Overall, higher LMA and leaf N distinguished the Chloridoideae/NAD‐ME group, while NADP‐ME and PCK grasses were distinguished by higher PNUE regardless of lineage. Plants were characterized by high photosynthesis and PWUE when grown at ambient CO₂ and by high conductance at inter‐glacial CO₂. In conclusion, the evolutionary and biochemical diversity among C₄ grasses was aligned with discernible leaf physiology, but it remains unknown whether these traits represent ecophysiological adaptation.     

Physiological advantages of C4 grasses in the field: a comparative experiment demonstrating the importance of drought

Global climate change is expected to shift regional rainfall patterns, influencing species distributions where they depend on water availability. Comparative studies have demonstrated that C₄ grasses inhabit drier habitats than C₃ relatives, but that both C₃ and C₄ photosynthesis are susceptible to drought. However, C₄ plants may show advantages in hydraulic performance in dry environments. We investigated the effects of seasonal variation in water availability on leaf physiology, using a common garden experiment in the Eastern Cape of South Africa to compare 12 locally occurring grass species from C₄ and C₃ sister lineages. Photosynthesis was always higher in the C₄ than C₃ grasses across every month, but the difference was not statistically significant during the wettest months. Surprisingly, stomatal conductance was typically lower in the C₃ than C₄ grasses, with the peak monthly average for C₃ species being similar to that of C₄ leaves. In water‐limited, rain‐fed plots, the photosynthesis of C₄ leaves was between 2.0 and 7.4 μmol m^?2 s^?1 higher, stomatal conductance almost double, and transpiration 60% higher than for C₃ plants. Although C₄ average instantaneous water‐use efficiencies were higher (2.4–8.1 mmol mol^?1) than C₃ averages (0.7–6.8 mmol mol^?1), differences were not as great as we expected and were statistically significant only as drought became established. Photosynthesis declined earlier during drought among C₃ than C₄ species, coincident with decreases in stomatal conductance and transpiration. Eventual decreases in photosynthesis among C₄ plants were linked with declining midday leaf water potentials. However, during the same phase of drought, C₃ species showed significant decreases in hydrodynamic gradients that suggested hydraulic failure. Thus, our results indicate that stomatal and hydraulic behaviour during drought enhances the differences in photosynthesis between C₄ and C₃ species. We suggest that these drought responses are important for understanding the advantages of C₄ photosynthesis under field conditions.     

Grass Physiognomic Trait Variation in African Herbaceous Biomes

African herbaceous biomes will likely face drastic changes in the near future, due to climate change and pressures from increasing human activities. However, these biomes have been simulated only by dynamic global vegetation models and failing to include the diversity of C₄ grasses has limited the accuracy of these models. Characterizing the floristic and physiognomic diversity of these herbaceous biomes would enhance the parameterization of C₄ grass plant functional types, thereby improving simulations. To this end, we used lowermost and uppermost values of three grass physiognomic traits (culm height, leaf length, and leaf width) available in most floras to identify several grass physiognomic groups that form the grass cover in Senegal. We then checked the capacity of these groups to discriminate herbaceous biomes and mean annual precipitation domains. Specifically, we assessed whether these groups were sufficiently generic and robust to be applied to neighboring (Chad) and distant (South Africa) phytogeographic areas. The proportions of two physiognomic groups, defined by their lowermost limits, delineate steppe from savanna and forest biomes in Senegal, and nama‐karoo, savanna, and grassland biomes in South Africa. Proportions of these two physiognomic groups additionally delineate the mean annual precipitation domains <600 mm and >600 mm in Senegal, Chad, and South Africa, as well as the <250 mm and >1000 mm domains in South Africa. These findings should help to identify and parameterize new C₄ grass plant functional types in vegetation models applied to West and South Africa.     

Ecological selection pressures for C4 photosynthesis in the grasses

Grasses using the C₄ photosynthetic pathway dominate grasslands and savannahs of warm regions, and account for half of the species in this ecologically and economically important plant family. The C₄ pathway increases the potential for high rates of photosynthesis, particularly at high irradiance, and raises water-use efficiency compared with the C₃ type. It is therefore classically viewed as an adaptation to open, arid conditions. Here, we test this adaptive hypothesis using the comparative method, analysing habitat data for 117 genera of grasses, representing 15 C₄ lineages. The evidence from our three complementary analyses is consistent with the hypothesis that evolutionary selection for C₄ photosynthesis requires open environments, but we find an equal likelihood of C₄ evolutionary origins in mesic, arid and saline habitats. However, once the pathway has arisen, evolutionary transitions into arid habitats occur at higher rates in C₄ than C₃ clades. Extant C₄ genera therefore occupy a wider range of drier habitats than their C₃ counterparts because the C₄ pathway represents a pre-adaptation to arid conditions. Our analyses warn against evolutionary inferences based solely upon the high occurrence of extant C₄ species in dry habitats, and provide a novel interpretation of this classic ecological association.     

Global patterns in local number of insect galling species

Abstract. We evaluate a three-part hypothesis explaining why gall-inducing insect species richness is so high in scleromorphic vegetation: (1) persistence of low nutrient status scleromorphic leaves facilitates the galling habit in warm temperate latitudes; (2) favourable colonization sites for gallers result from reduced hygrothermal stress, high phenolics in the outer cortex of the gall, and reduced carnivore and fungal attack in the gall; and (3) in more mesic sites, mortality is high due to carnivore attack and invasion of galls by fungi. Over 280 samples of local species of galling herbivorous insects from fourteen countries on all continents except Antarctica revealed a strong pattern of highest richness in warm temperate latitudes, or their altitudinal equivalents. The peak of galling species richness on the latitudinal gradient from the equator into the Arctic was between 25 to 38° N or S. Galling species were particularly diverse in sclerophyllous vegetation, which commonly had greater than twelve species per local sample. In mesic, non-sclerophyllous vegetation types the number of galling species was lower with twelve or fewer species present. Many sites in sclerophyllous vegetation supported between thirteen and forty-six galling species locally, including campina islands in Amazonia, cerrado savanna in central Brazil, the Sonoran Desert in Arizona and Mexico, shrubland in Israel, fynbos in South Africa and coastal scleromorphic vegetation in Australia. At the same latitude, or its elevational equivalent, galling species richness was significantly higher in relatively xeric sites when compared to riparian or otherwise mesic habitats, even when scleromorphic vegetation dominated the mesic sites. The results were consistent with the hypothesis and extend to a more general level the patterns and predictions on the biogeography of gall-inducing insects.     

A revised evolutionary history of Poales: origins and diversification

Poales represents more than one‐third of all monocotyledons (c. 20 000 species in 16 families) and constitutes a microcosm of the angiosperms. The extreme variation in species richness among the families of Poales is still not understood: Poaceae includes ～10 000 species, whereas six families have fewer than ten species. Here, using the largest phylogenetic analysis of Poales to date, molecular dating, ancestral reconstructions and diversification analyses, we develop a macro‐evolutionary and macro‐ecological approach to seek correlates for changing diversification patterns. We show that the poalean families diverged in the Late Cretaceous, a time of high levels of CO₂ and high rainfall. Our habitat reconstructions indicate that Poales inhabited open and dry habitats in this environment. We also demonstrate that lineages with CO₂‐concentrating mechanisms inhabiting dry and open environments exhibited higher diversification rates than C₃, shade and wet lineages. CO₂‐concentrating mechanisms counteract the effects of low atmospheric CO₂ and reduce phototranspiration. It is believed that the parallel evolution of C₄ and CAM (Crassulacean acid metabolism) photosynthesis in Poaceae, Cyperaceae and Bromeliaceae is an adaptation to changes in atmospheric CO₂ concentrations. Combinations of extrinsic and intrinsic factors might have played a role in shifts in diversification rates and may explain the variation in species richness in Poales. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 4–16.