Aboveground Biomass Removal by Burning and Raking Increases Diversity in a Reconstructed Prairie

Prescribed spring burning often contributes to a predominance of C₄ grasses and low forb abundance and is impractical at many sites, especially near development. We tested raking after mowing as an alternative to prescribed burning in a reconstructed Minnesota prairie. We also tested mowing without raking as a possible means of maintaining prairie communities. Frequency, flowering stem abundance, and cover were measured for all plant species and native functional groups (C₄ grasses, C₃ graminoids, forbs, legumes, and annual or biennial forbs). Mowing alone did not differ from the control in its effect on any functional groups of plants. Round‐headed bush clover (Lespedeza capitata), a legume, and Black‐eyed Susan (Rudbeckia hirta), a biennial, increased in frequency with treatments that removed biomass (i.e., fire or raking), but they did not have significantly more flowering stems. Thus, new plants established well from seed, whereas the vitality of mature plants did not change. Raking had similar effects to burning on most functional groups, although flowering stems of C₄ grasses were significantly more abundant after fire than after raking. Burning reduced some C₃ forbs and grasses and favored the dominance of C₄ grasses. Therefore, raking after mowing in the spring provides an alternative to prescribed burning that has many of the same positive aspects as fire but does not promote aggressive C₄ grasses to the same extent.     

Sulcal patterns of fossilTheropithecus baboons: Phylogenetic and functional implications

Sulcal patterns were determined from endocasts of fossilTheropithecus (T. oswaldi andT. darti) and compared to the sulcal pattern of extant geladas (T. gelada). A comparison of the configurations of central and arcuate sulci suggests that the cortical motor face representation is relatively enlarged in extantTheropithecus, whereas the cortical motor and sensory face representations were enlarged inT. oswaldi. A relatively expanded sensory and/or motor cortical face representation may be related to masticatory (e.g., “seed-eating”) and gestural (e.g., “lipflip”) functions. SinceT. oswaidi is more derived (expanded) in sensory face representation thanT. gelada, it seems unlikely that it was a direct ancestor of modern geladas. Rather, the two groups were separate by 1.8 million years ago and may have had a most recent common ancestor similar toT. darti from the Hadar Formation of Ethiopia between 2 and 3 million years ago.     

Summer feeding strategy of Spanish ibexCapra pyrenaica and domestic sheepOvis aries in south-eastern Spain

The feeding strategies of the Spanish ibexCapra pyrenaica Schinz, 1838 and sheepOvis aries Linnaeus, 1758 during summer in the alpine area of Sierra Nevada in south-eastern Spain were compared. The degree of vegetation use by the two ungulates, food availability, diet composition, species selection indices and the effect of plant availability and chemical composition on diet selection were analysed. The available biomass of herbaceous and woody resources was 87 and 13%, respectively. The degree of use was 15% for herbaceous vegetation and 10.3% for woody vegetation. The most used species were those with moderate amounts in diet composition of both ruminants. The main plant groups consumed by Spanish ibex were graminoids (57.5%), shrubs (20.5%) and chamaephytes (14.3%). Sheep consumed graminoids (41.6%), forbs (32.3%) and chamaephytes (20.9%). There was a significant correlation between the two diets, with an overlap of 56.3%. The highest indices of selection were found in items with moderate-low availability and which comprised moderate levels in the diet composition of the two species. Spanish ibex mainly selected diet according to availability, while sheep selection depended on protein content. The ibex consumed cell wall-rich resources such as grasses and shrubs, while the sheep consumed higher quality feed and includes more forbs in its diet.     

Assessing Targets for the Restoration of Herbaceous Vegetation in Ponderosa Pine Forests

A restoration project is considered a success when the initial target is met, but many targets are plausible. We evaluated the success of a restoration project in its 11th year since treatment in a southwestern ponderosa pine–bunchgrass community and the appropriateness of several targets. We measured the responses of (1) total standing crop; (2) standing crop of five functional groups (C₃ and C₄ graminoids, leguminous forbs, and nonleguminous perennial and annual forbs); (3) graminoid community composition; and (4) standing crop of five common graminoid species (Festuca arizonica, Muhlenbergia montana, Elymus elymoides, Carex geophila, and Poa fendleriana). Targets were quantified in remnant grass patches, which provided the standards for these targets, and were assessed in three other forest patch types (pre‐settlement tree patches, post‐settlement tree patches, and patches where all post‐settlement trees were removed). Patches where all post‐settlement trees were removed reached target levels for total standing crop, C₃ and C₄ graminoid standing crop, graminoid community composition, and M. montana, E. elymoides, and C. geophila standing crops. Standing crop of legumes and of F. arizonica did not increase over time in any patch type. Targets were not met in pre‐settlement patches or in patches where some post‐settlement trees were left standing, suggesting that it is unrealistic to expect equal responses across all patch types. If increasing herbaceous standing crop is a major goal, practitioners should create gaps within the pine forest canopy.     

Stable isotopic analysis of human diet in the Marianas Archipelago,Western Pacific

Proportions of marine vs. terrestrial resources in prehistoric human diets in the southern Mariana Islands (Guam, Rota, Saipan), Micronesia, have been estimated by analysis of stable isotope ratios of carbon and nitrogen in bone collagen and of carbon in apatite. The isotopic composition of marine and terrestrial food resources from the Marianas have also been determined. Experimental evidence shows that collagen carbon isotopes mainly reflect those of dietary protein sources and thus overestimate the contribution of marine animal foods. Marine protein consumption apparently ranges from ∼20% to ∼50% on these islands. Experiments also demonstrate the carbon isotope ratio of bone apatite carbonate accurately reflects that of the whole diet. Carbonate carbon isotope data suggest some individuals consumed significant amounts of ¹³C-enriched (C₄) plants or seaweeds. Sugar cane is an indigenous C₄ crop and seaweeds are eaten throughout the Pacific, but they have not been considered by archaeologists to have been prehistoric dietary staples. Apatite carbon isotope analysis has apparently identified previously unrecognized prehistoric dietary adaptations in the Mariana Islands, but this must be confirmed by archaeobotanical evidence. Am J Phys Anthropol 104:343–361, 1997. © 1997 Wiley-Liss, Inc.     

Invasive forbs differ functionally from native graminoids, but are similar to native forbs

? Exotic plant invasions can alter ecosystem processes, particularly if the invasive species are functionally different from native species. We investigated whether such alterations can be explained by differences in functional traits between native and invasive plants of the same functional group or by differences in functional group affiliation. ? We compared six invasive forbs in Europe with six native forbs and six native graminoids in leaf and whole-plant traits, plasticity in response to nutrient supply and interspecific competition, litter decomposition rate, effects on soil nutrient availability, and allelopathy. All traits were measured in a series of pot experiments, and leaf traits additionally in the field. ? Invasive forbs differed from native forbs for only a few traits; they had less leaf chlorophyll and lower phosphorus (P) uptake from soil, but they tended to have a stronger allelopathic effect. The invasive forbs differed in many traits from the native graminoids, their leaves had lower tissue densities and a shorter life span, their litter decomposed faster and they had a lower nitrogen-use efficiency. ? Our results suggest that invasive forbs have the potential to alter ecosystem properties when invading graminoid-dominated and displacing native graminoids but not when displacing native forbs.     

Predicting plant response to arbuscular mycorrhizas: The role of host functional traits

The arbuscular mycorrhiza (AM) is among the most ubiquitous symbiosis in the world. A meta-analysis of 759 articles (1978–2012) was conducted to test whether ecologically important host plant traits (N-fixation and C-fixation pathway) affect the response of the plant to mycorrhizal colonization. We found that the effect of N-fixation on mycorrhizal growth response (MGR) depended on whether the plant was woody or a forb. N-fixing forbs had a higher MGR than non-N-fixing forbs, but the reverse was true for woody plants. Moreover, C₄-grasses had significantly higher MGR than C₃-grasses, but no significant difference was found between C₃ and C₄ forbs, or between C₃ and C₄ woody species. Overall, woody species had higher MGR than any other functional group. These results demonstrate that MGR does depend on host functional characteristics, but neither N-fixation capacity nor C-fixation pathway are apparently fundamental controllers of MGR. Instead, it would appear possible that these traits influence MGR only insofar as they influence more fundamental functions such as P demand and P supply.     

Changes in the Herbaceous Communities on the Landslide of the Casita Volcano, Nicaragua, during Early Succession

This study examined the main changes in the herbaceous communities during the first four years of succession in a large landslide on Casita Volcano, Nicaragua, located in a densely populated area that has a tropical dry climate. Our main objective was to determine the major pathways of change in community features, such as richness, biovolume, species composition, and abundances of plant traits and to verify if they varied between the different landslide areas based on abiotic heterogeneity and landscape context. Number, percent cover, mean height and traits of herbaceous species, and several abiotic factors related to fertility and geomorphological stability of substrates were sampled in 28 permanent plots. Environmental heterogeneity strongly influenced early successional changes in the herbaceous communities during the four years of the study. Biovolume increased in the unstable and infertile areas and decreased in the stable and fertile landslide areas. In most zones, species richness decreased significantly because of the expansion of a few dominant species that developed a large cover and excluded other species. Those dominant species were mainly responsible for changes in species composition and included annual forbs, e.g., Calopogonium mucunoides and Stizolobium pruriens, graminoids that have rhizomes or stolons, e.g., Hyparrhenia rufa, and perennial nitrogen-fixing forbs, e.g., Clitoria ternatea. They might be important in determining future successional patterns on the Casita Volcano landslide.     

Isotopic reconstructions of habitat change surrounding the extinction of Sivapithecus,a Miocene hominoid,in the Siwalik Group of Pakistan

This research presents an isotopic study of a wide range of mammalian taxa from the Miocene Siwalik Group of Pakistan, with a focus on two time intervals—9.3–9.2 Ma, when the hominoid Sivapithecus was present, and 8.1–8.0 Ma, shortly after Sivapithecus became extinct. The purpose of this investigation is to reconstruct the vegetation mosaic at both time levels in order to determine what Sivapithecus habitat was like and what changes in habitat and climate may have taken place by the time Sivapithecus went extinct. Both carbon and oxygen stable isotopes were sampled from inorganic carbonate in tooth enamel. Results indicate a vegetation mosaic of both closed and open habitat at both time intervals, but a decrease in forest accompanied by an increase in open habitat (including C₄ grasses) during the younger interval. Individuals from many species cluster with each other with respect to both carbon and oxygen isotope values, indicating a spectrum of feeding adaptations exploiting different parts of the habitat. Isotopic values for Sivapithecus suggest that it fed in the forest upper canopy. Taxa feeding in the most closed, wet habitat become extinct by 8.1 Ma. Furthermore, higher δ¹⁸O values over time suggest a change in climate with a reduction in annual rainfall, perhaps accompanied by changes in precipitation sources or rainfall regime as well. These results suggest that forests became fragmented over time. While still present in the younger level, much of the forest was replaced by open habitat, including patches of C₄ grass. Forest loss and fragmentation is a likely cause of the extinction of Sivapithecus.     

Changes in plant form and function across altitudinal and wetness gradients in the wetlands of the Maloti-Drakensberg,South Africa

A survey of 93 wetlands in six catchments across the Maloti-Drakensberg is used to assess the distribution of plant functional types across altitudinal and wetness gradients. Altitudes range from 1,000 to 3,200 m a.s.l. Within each catchment, the wetlands were selected to cover the complete range in altitude and wetland types. In each of the selected wetlands, vegetation was sampled in 3 by 3 m quadrats covering the entire range of wetness represented in the wetland, from temporarily wet to permanently inundated soils. Plant species were allocated to one of 11 different functional types (examples are C₃ grasses, C₄ sedges, rosette plants, and shrubs), and the proportion of the vegetation in each sample occupied by each functional type was calculated from the species’ abundances. Canonical Correspondence Analysis shows that “wetness” clearly has the highest impact on the distribution of functional types, followed by altitude. The most important plant functional types in wetlands are grasses and sedges, however, at higher altitudes, forbs (especially rosette plants) and bulbous plants become a more prominent feature in the wetlands. The total amount of graminoids gradually decreases with altitude. The general trend is that sedges tend to increase with increasing wetness and C₃ plants (grasses and sedges) increase with increasing altitude, but these effects are not independent. The distributions of C₄ sedges and C₄ grasses along an altitudinal gradient are quite different, and C₄ grasses grow abundantly at much higher altitudes than C₄ sedges. C₄ sedges are very scarce at the altitudes represented in the Maloti-Drakensberg area, whereas C₃ grasses occur in the permanently wet parts of the wetlands, especially at higher altitudes (normally mostly occupied by sedges). Shrubs are rare in wetlands and tend to be an indication of disturbance. This study complements previous studies on the distribution of grasses and sedges at the lower altitudes within KwaZulu-Natal, which found that at altitudes below 1,000 m a.s.l. C₄ sedges were much more prominent, while forbs and rosette plants were largely absent. This confirms that C₄ as an adaptation to hotter and warmer climates is sometimes a less favorable metabolism in wet high altitude areas. At high altitudes, rosette plants and bulbous plants become more competitive in wetlands, probably because grasses and sedges present at these altitudes generally grow smaller than they do in low altitude wetlands.     

Assessing the nutritional consequences of switching foraging behavior in wood bison

Diet is one of the most common traits used to organize species of animals into niches. For ruminant herbivores, the breadth and uniqueness of their dietary niche are placed on a spectrum from browsers that consume woody (i.e., browse) and herbaceous (i.e., forbs) plants, to grazers with graminoid‐rich diets. However, seasonal changes in plant availability and quality can lead to switching of their dietary niche, even within species. In this study, we examined whether a population of wood bison (Bison bison athabascae) in northeast Alberta, Canada, seasonally switched their foraging behavior, and if so, whether this was associated with changes in nutrient acquisition. We hypothesized that bison should switch foraging behaviors from grazing in the winter when standing, dead graminoids are the only foliar plants readily available to browsing during spring and summer as nutritious and digestible foliar parts of browse and forbs become available. If bison are switching foraging strategy to maximize protein consumption, then there should be a corresponding shift in the nutritional niche. Alternatively, if bison are eating different plants, but consuming similar amounts of nutrients, then bison are switching their dietary niche to maintain a particular nutrient composition. We found wood bison were grazers in the winter and spring, but switch to a browsing during summer. However, only winter nutrient consumption of consumed plants differed significantly among seasons. Between spring and summer, bison maintained a specific nutritional composition in their diet despite compositional differences in the consumed plants. Our evidence suggests that bison are selecting plants to maintain a target macronutrient composition. We posit that herbivore''s can and will switch their dietary niche to maintain a target nutrient composition.     

Photosynthetic and morphological functional types for native species from mixed prairie in Southern Saskatchewan, Canada

Photosynthetic pathways (e.g. C₃, C₄) and morphological functional types (e.g. trees, shrubs, high perennial grasses, perennial forbs) were identified for the native species from the Saskatchewan mixed prairie, using the data from references published between 1950 and 2003. Of the total 219 identified species in 145 genera and 45 families, 208 species in 137 genera and 44 families were found with C₃ photosynthesis, and most of these species are dominants (e.g. Agropyron dasystachyum Hook. and Stipa spartea var. curtiseta Hitchc.). 11 species in 10 genera and 3 families were identified with C₄ photosynthesis (e.g. Atriplex argentea Nutt., Andropogon scoparius Michx., Boutelou gracilis Lag., Calamovilfa longifolia Hook.). The amount of total identified C₄ species in the region is much less than that from the South Dakota mixed prairie (27 species). Gramineae is the leading family with C₄ photosynthesis (8 species), Chenopodiaceae ranks the second (2 species). Relatively less forb types [50 % perennial forbs (PEF) and 12 % annual forbs (ANF)] and more graminoid types (25 %) composition suggested that the rangelands in the region are relatively stable. Lacking of the knowledge on the optimal traits for PFTs classification in the region, further studies (e.g. C₃ and C₄ plant identification and optimal trait selection) are needed to explore the relationships between PFTs and vegetation variations, as well as land-use and climate changes.     

Effects of climate and atmospheric CO2 partial pressure on the global distribution of C4 grasses: present, past, and future

C₄ photosynthetic physiologies exhibit fundamentally different responses to temperature and atmospheric CO₂ partial pressures (pCO₂) compared to the evolutionarily more primitive C₃ type. All else being equal, C₄ plants tend to be favored over C₃ plants in warm humid climates and, conversely, C₃ plants tend to be favored over C₄ plants in cool climates. Empirical observations supported by a photosynthesis model predict the existence of a climatological crossover temperature above which C₄ species have a carbon gain advantage and below which C₃ species are favored. Model calculations and analysis of current plant distribution suggest that this pCO₂-dependent crossover temperature is approximated by a mean temperature of 22°C for the warmest month at the current pCO₂ (35 Pa). In addition to favorable temperatures, C₄ plants require sufficient precipitation during the warm growing season. C₄ plants which are predominantly graminoids of short stature can be competitively excluded by trees (nearly all C₃ plants) – regardless of the photosynthetic superiority of the C₄ pathway – in regions otherwise favorable for C₄. To construct global maps of the distribution of C₄ grasses for current, past and future climate scenarios, we make use of climatological data sets which provide estimates of the mean monthly temperature to classify the globe into areas which should favor C₄ photosynthesis during at least 1 month of the year. This area is further screened by excluding areas where precipitation is <25 mm per month during the warm season and by selecting areas classified as grasslands (i.e., excluding areas dominated by woody vegetation) according to a global vegetation map. Using this approach, grasslands of the world are designated as C₃, C₄, and mixed under current climate and pCO₂. Published floristic studies were used to test the accuracy of these predictions in many regions of the world, and agreement with observations was generally good. We then make use of this protocol to examine changes in the global abundance of C₄ grasses in the past and the future using plausible estimates for the climates and pCO₂. When pCO₂ is lowered to pre-industrial levels, C₄ grasses expanded their range into large areas now classified as C₃ grasslands, especially in North America and Eurasia. During the last glacial maximum (∼18 ka BP) when the climate was cooler and pCO₂ was about 20 Pa, our analysis predicts substantial expansion of C₄ vegetation – particularly in Asia, despite cooler temperatures. Continued use of fossil fuels is expected to result in double the current pCO₂ by sometime in the next century, with some associated climate warming. Our analysis predicts a substantial reduction in the area of C₄ grasses under these conditions. These reductions from the past and into the future are based on greater stimulation of C₃ photosynthetic efficiency by higher pCO₂ than inhibition by higher temperatures. The predictions are testable through large-scale controlled growth studies and analysis of stable isotopes and other data from regions where large changes are predicted to have occurred. Received: 3 July 1997 / Accepted: 3 December 1997     

Foliar macronutrient concentrations of forest understorey species in relation to Ellenberg's indices and potential relative growth rate

In order to investigate broad patterns of variation of the foliar mineral nutrient concentrations of herbaceous plant communities in the ground layer of W Europe forests, correlations were examined between Ellenberg's indices (N-index: mineral nitrogen availability, R-index: pH, F-index: soil moisture and L-index: light intensity) and literature values of macronutrient concentrations for 84 forbs and 39 graminoid species. Significant, positive correlations were found between the R-index and the plants' concentration of Ca and K (forbs only) and between the N-index and the plants' concentration of K, P (forbs only) and N (forbs and graminoids). Multiple regressions showed that the N-index was the best predictor of the plants' concentration of N (forbs and graminoids), P (forbs) and Ca (graminoids) and the R-index of the plants' concentration of Ca and K (forbs). The mineral nutrient concentrations of graminoids were lower and less strongly correlated with Ellenberg's indices than those of forbs. It is argued that the mineral nutrient concentrations in the plants match the availability of mineral nutrients in the soil for N, P (N-index) and Ca (R-index), but not for K and Mg. Significant, positive correlations were found between potential relative growth rate and the concentration of some elements (N, P, K and Ca in forbs, Ca and Mg in graminoids). This suggests that the increase in the concentration of these elements in plants along fertility gradients is due, at least partly, to genetically controlled alterations of leaf anatomy associated with increasing potential relative growth rate.     

Carbon isotope ratios and crop analyses of Arphia (Orthoptera: Acrididae) species in southeastern Wyoming Grassland

Summary Food habits of Arphia conspersa Scudder and Arphia pseudonietana (Thomas) were studied along an altitudinal transect in southeastern Wyoming shortgrass mixed prairie. Stable carbon isotope ratios indicated that diets were significantly different between study sites, between species, and between sexes. These differences were found to be primarily related to the availability of different food plants along the transect, although species with the C₃ pathway of photosynthesis were consumed in greater proportion than their availability in the habitat. The preference for C₃ species is presumably related to their higher nutritional value and digestibility, in spite of the fact that more time and energy must be spent to locate these food plants in some of the habitats studied. This study demonstrates the utility of the carbon isotope method in studying plant-animal interactions in habitats containing both C₃ and C₄ plants.     

Caratteri Fondamentali della Vegetazione delle Paludi Pontine

Abstract

Observations in various microhabitats as regards the composition of different growth forms in an alpine region of Tungnath, northwestern Himalaya, were carried out. Maximum frequency was displayed by mat-forming forbs (Trachydium roylei, 100%) throughout the growing season (May–October). Maximum values for density and Importance Value Index (IVI) (452.10 plants m^?2 and 157.8, respectively) were recorded for tussock graminoids, Poa annua (L.). The IVI of different growth forms exhibited significant positive correlation with stoloniferous graminoids (KD), mat forming forbs (TR) and prostrate creeping dwarf shrubs (GT). A significant positive correlation with plant available water was also detected for mat-forming forbs (TR) and one of the plant species (CN) representing stoloniferous graminoids. The values of concentration of dominance (Cd) ranged from 0.0001 to 0.277. Diversity index (H) was highest (0.532) for tussock graminoids [Danthonia cachemyriana (Jaub &; Spach.), P. annua] indicating the relative abundance of this growth form. Overall, tussock graminoids are dominant in the region, indicating stable and moderate steepness of the slopes, the major topographic feature of Tungnath. Besides topography, grazing, anthropogenic activities and interactions among associated species are the determining factors for the phytostructure of alpine communities in Himalaya.     

Patterns of water potential and photosynthesis of desert sand dune plants,Eureka Valley,California

Summary This study examined the mode of photosynthesis (C₃ or C₄), daily and seasonal patterns of xylem water potential, seasonal patterns of field photosynthesis, and the laboratory gas exchange characterisitcs of plants which grow on or in the vicinity of Eureka Dunes, Inyo County, California. The perennial duneendemic Swallenia alexandrae was found to possess the C₄ pathway while all other taxa surveyed were C₃. Plants which grew on the dunes exhibited: 1) significantly less negative xylem water potentials, 2) dampened seasonal changes in predawn water potentials, and 3) smaller seasonal amplitudes of water potential than plants of the adjoining flats. The minimum water potentials experienced by Swallenia during the hot summer months were a third of those endured by adjacent non-dune Larrea. Non-endemics growing on the dune had more negative xylem water potentals than dune endemics, but still never approached the low values of non-dune plants. The poor moisture retention properties of sand may have selected for moisture-conserving traits (stomatal closure at relatively high water potentials, high water use efficiency) rather than moisture-expending ones (osmoregulation, high leaf conductances) in the endemic perennials. Field measurements of photosynthesis showed that dune-restricted (but not necessarily endemic) plants had high photosynthetic capacities and sustained summer carbon assimilation, the latter being protracted months beyond the last pulse of precipitation. The C₃ annual Dicoria canescens ssp. clarkae maintained photosynthetic rates well exceeding those of the C₄ Swallenia throughout the summer and may represent a previously undescribed physiological life form in desert plants. Laboratory measurements supplemented the field data and compared the water use efficiencies of two dune endemics. It is suggested that high photosynthetic productivity, high water use efficiency, and carbon allocation to the longitudinal growth of roots and shoots are important physiological adaptations to shifting sand and substrate moisture depletion at Eureka Dunes.     

Environmental and Evolutionary Preconditionsfor the Origin and Diversification of the C4 PhotosyntheticSyndrome

Abstract: C₄ photosynthesis is an evolutionary solution to high rates of photorespiration and low kinetic efficiency of Rubisco in CO₂‐depleted atmospheres of recent geologic time. About 7500 plant species are C₄, in contrast to 30 000 CAM and 250 000 C₃ species. All C₄ plants occur in approximately 90 genera from 18 angiosperm families. In all of these families, the C₄ pathway evolved independently. In many, multiple independent origins have occurred, such that over 30 distinct evolutionary origins of the C₄ pathway are recognized. Fossil and carbon isotope evidence show that the C₄ syndrome is at least 12 to 15 million years old, although estimates based on molecular sequence comparisons indicate it is over 20 million years old. The evolutionary radiation of herbaceous angiosperms may have been required for C₄ plant evolution. All C₄ species occur in advanced angiosperm families that appeared in the fossil record in the past 70 million years. Most of these families diversified in terms of genera and species numbers between 20 to 40 million years ago, during a period of global cooling, atmospheric CO₂ reduction and aridification. During the period of diversification, numerous traits arose in the C₃ flora that enhanced their performance in arid environments and atmospheres of reduced CO₂. Some of these traits may have predisposed certain taxa to develop the C₄ pathway once atmospheric CO₂ levels declined to a point where the ability to concentrate CO₂ had a selective advantage. Leading traits in C₃ plants that may have facilitated the initial transition to C₄ photosynthesis include close vein spacing and an enlargement of the bundle sheath cell layer to form a Kranz‐like anatomy. Ecological factors not directly connected with photosynthesis probably also played a role. For example, extensive ecological disturbance may have been needed to convert C₃‐dominated woodlands into open, high‐light habitats where herbaceous C₄ plants could succeed. Disturbances in the form of fire, and browsing by large mammals, increase during the time of C₄ plant evolution and diversification. Fire increased because of the drying climate, while browsing increased with the evolutionary diversification of the mammalian megafauna in the Oligocene and Miocene epochs. In summary, the origin of C₄ plants is hypothesized to have resulted from a novel combination of environmental and phylogenetic developments that, for the first time, established the preconditions required for C₄ plant evolution.     

Browsing and grazing in elephants: the isotope record of modern and fossil proboscideans

The diet of extant elephants (Loxodonta in Africa, Elephas in Asia) is dominated by C₃ browse although some elephants have a significant C₄ grass component in their diet. This is particularly noteworthy because high-crowned elephantid cheek teeth represent adaptation to an abrasive grazing diet and because isotopic analysis demonstrates that C₄ vegetation was the dominant diet for Elephas in Asia from 5 to 1 Ma and for both Loxodonta and Elephas in Africa between 5–1 Ma. Other proboscideans in Africa and southern Asia, except deinotheres, also had a C₄-dominated diet from about 7 Ma (when the C₄ biomass radiated in tropical and subtropical regions) until their subsequent extinction. Received: 1 July 1998 / Accepted: 16 February 1999     

Granivory reduces biomass and lignin concentrations of plant tissue during grassland assembly

Small mammals can influence grassland assembly by selecting against palatable plants – the community can become dominated by the plants they avoid. This predation-based selection could have indirect effects on community biomass and tissue quality, especially given how untasty plants may have higher concentrations of recalcitrant carbon compounds including lignin. We tested small mammal effects on biomass and tissue quality of roots and shoots in a two-year-old 18 ha restored tallgrass prairie with established zones of high and low plant predation. We focused on the three dominant herbaceous functional groups of tallgrass prairie (perennial forbs, C₃ and C₄ grasses), and targeted the early stages of assembly given that plant predation by small animals can unfold quickly and is difficult to subsequently quantify. We predicted rodent predation to create communities with reduced biomass but an increased abundance of lignin-rich plants; we only observed the former. Rodents reduced aboveground biomass by 46% but preferentially targeted lignin-rich plants, with the latter result explained by the predominance of granivory over herbivory – there was no opportunity for selection based on tissue palatability. Based strictly on aboveground biomass, we estimated small mammals reduced standing stocks of recalcitrant carbon by 65 kg ha⁻¹, with reductions in belowground stocks almost certainly higher given that root:shoot ratios averaged 21:1. Given that the quantity and quality of plant production can affect ecosystem functions including decomposition and the regulation of soil carbon stocks, our work suggests that non-random plant predation may substantially affect rates of soil carbon accumulation in the early stages of grassland development.