Fire and the Miocene expansion of C4 grasslands

C₄ photosynthesis had a mid‐Tertiary origin that was tied to declining atmospheric CO₂, but C₄‐dominated grasslands did not appear until late Tertiary. According to the ‘CO₂‐threshold’ model, these C₄ grasslands owe their origin to a further late Miocene decline in CO₂ that gave C₄ grasses a photosynthetic advantage. This model is most appropriate for explaining replacement of C₃ grasslands by C₄ grasslands, however, fossil evidence shows C₄ grasslands replaced woodlands. An additional weakness in the threshold model is that recent estimates do not support a late Miocene drop in pCO₂. We hypothesize that late Miocene climate changes created a fire climate capable of replacing woodlands with C₄ grasslands. Critical elements were seasonality that sustained high biomass production part of year, followed by a dry season that greatly reduced fuel moisture, coupled with a monsoon climate that generated abundant lightning‐igniting fires. As woodlands became more open from burning, the high light conditions favoured C₄ grasses over C₃ grasses, and in a feedback process, the elevated productivity of C₄ grasses increased highly combustible fuel loads that further increased fire activity. This hypothesis is supported by paleosol data that indicate the late Miocene expansion of C₄ grasslands was the result of grassland expansion into more mesic environments and by charcoal sediment profiles that parallel the late Miocene expansion of C₄ grasslands. Many contemporary C₄ grasslands are fire dependent and are invaded by woodlands upon cessation of burning. Thus, we maintain that the factors driving the late Miocene expansion of C₄ were the same as those responsible for maintenance of C₄ grasslands today.     

Elevation‐induced climate change as a dominant factor causing the late Miocene C4 plant expansion in the Himalayan foreland

During the late Miocene, a dramatic global expansion of C₄ plant distribution occurred with broad spatial and temporal variations. Although the event is well documented, whether subsequent expansions were caused by a decreased atmospheric CO₂ concentration or climate change is a contentious issue. In this study, we used an improved inverse vegetation modeling approach that accounts for the physiological responses of C₃ and C₄ plants to quantitatively reconstruct the paleoclimate in the Siwalik of Nepal based on pollen and carbon isotope data. We also studied the sensitivity of the C₃ and C₄ plants to changes in the climate and the atmospheric CO₂ concentration. We suggest that the expansion of the C₄ plant distribution during the late Miocene may have been primarily triggered by regional aridification and temperature increases. The expansion was unlikely caused by reduced CO₂ levels alone. Our findings suggest that this abrupt ecological shift mainly resulted from climate changes related to the decreased elevation of the Himalayan foreland.     

C4 photosynthesis in the vegetation of Aldabra Atoll

Summary The occurrence of the C₄ photosynthetic system, based upon an examination of leaf anatomy, of nearly all species of the vegetation of Aldabra Atoll, western Indian Ocean, is reported. About 19% of the flora has the C₄ system. It only occurs in herbaceous plants, which grow either in grasslands or as an understorey to tall shrubs. Both C₃ and C₄ species compete side by side in the mixed-species communities.The Royal Society Aldabra Research Station     

Oligocene CO2 decline promoted C4 photosynthesis in grasses

C4 photosynthesis is an adaptation derived from the more common C3 photosynthetic pathway that confers a higher productivity under warm temperature and low atmospheric CO2 concentration [1, 2]. C4 evolution has been seen as a consequence of past atmospheric CO2 decline, such as the abrupt CO2 fall 32-25 million years ago (Mya) [3-6]. This relationship has never been tested rigorously, mainly because of a lack of accurate estimates of divergence times for the different C4 lineages [3]. In this study, we inferred a large phylogenetic tree for the grass family and estimated, through Bayesian molecular dating, the ages of the 17 to 18 independent grass C4 lineages. The first transition from C3 to C4 photosynthesis occurred in the Chloridoideae subfamily, 32.0-25.0 Mya. The link between CO2 decrease and transition to C4 photosynthesis was tested by a novel maximum likelihood approach. We showed that the model incorporating the atmospheric CO2 levels was significantly better than the null model, supporting the importance of CO2 decline on C4 photosynthesis evolvability. This finding is relevant for understanding the origin of C4 photosynthesis in grasses, which is one of the most successful ecological and evolutionary innovations in plant history.     

C4 Photosynthesis Promoted Species Diversification during the Miocene Grassland Expansion

Identifying how organismal attributes and environmental change affect lineage diversification is essential to our understanding of biodiversity. With the largest phylogeny yet compiled for grasses, we present an example of a key physiological innovation that promoted high diversification rates. C₄ photosynthesis, a complex suite of traits that improves photosynthetic efficiency under conditions of drought, high temperatures, and low atmospheric CO₂, has evolved repeatedly in one lineage of grasses and was consistently associated with elevated diversification rates. In most cases there was a significant lag time between the origin of the pathway and subsequent radiations, suggesting that the ‘C₄ effect’ is complex and derives from the interplay of the C₄ syndrome with other factors. We also identified comparable radiations occurring during the same time period in C₃ Pooid grasses, a diverse, cold-adapted grassland lineage that has never evolved C₄ photosynthesis. The mid to late Miocene was an especially important period of both C₃ and C₄ grass diversification, coincident with the global development of extensive, open biomes in both warm and cool climates. As is likely true for most “key innovations”, the C₄ effect is context dependent and only relevant within a particular organismal background and when particular ecological opportunities became available.     

Faunal change in the Turkana Basin during the late Oligocene and Miocene

Faunal evolution over the last 65 million years of earth's history was dominated by mammalian radiations, but much of this era is poorly represented in Africa. Mammals first appeared early in the Mesozoic, living alongside dinosaurs for millions of years, but it was not until the extinction of dinosaurs 65 myr ago that the first major explosion of mammalian taxa took place. The Cenozoic (65 Ma to Recent) witnessed repeated and dynamic events involving the radiation, evolution, and extinction of mammalian faunas. Two of these events, each marking the extinction of one diverse fauna and subsequent establishment of another equally diverse fauna, both involving advanced catarrhine primates, are recorded in sites in the Turkana Basin, despite the poorly represented record of Cenozoic faunas elsewhere in sub-Saharan Africa. The first of these events occurred at the Oligocene-Miocene transition and the other at the Miocene-Pliocene transition.     

Zonation of late Miocene and early Pliocenecircum-mediterranean faunas

This attempt at biostratigraphic correlations for the fossiliferous sites of the late Miocene and early Pliocene,situated around the Mediterranean basin, is based on the zones established by P. Mein according to variations in the mammalian faunas. New taxa are mentioned, particularly carnivores; moreover, a series of African localities are integrated and, for some of them, an analysis of the faunal constituents leads to paleobiogeographic considerations.     

Early Miocene vegetation and climate in Weichang District, North China

The Early Miocene palynological assemblage of Guangfayong (GFY) in the Weichang District, Hebei Province, China has been studied. It consists of 48 palynomorphs belonging to 39 families, with pollen and spores belonging to angiosperms (28.9%), gymnosperms (59.9%), ferns (10.8%) and other elements (0.5%). Based on the palynological assemblages of GFY and Wuluogong (WLG), another locality in the Weichang District, the Early Miocene vegetation of the Weichang District, was characterized by a mixed temperate forest of conifers (e.g. Pinus, Picea, Tsuga) and broad-leaved trees (e.g. Betula, Alnus), with some subtropical plants (e.g. Carya). The palaeoclimatic parameters of Guangfayong were obtained by applying the Coexistence Approach: the mean annual temperature from 7.8 to 14.9 °C, the difference of temperature between the coldest and warmest months from 14.2 to 23 °C, the mean temperature of the coldest month from − 3 to 5.9 °C, the mean temperature of the warmest month from 23.5 to 25.4 °C, the mean annual precipitation from 658.7 to 1389.4 mm, the minimum monthly precipitation from 7.6 to 16.4 mm, and the maximum monthly precipitation from 161.4 to 205.9 mm. It suggests a warm temperate to subtropical climate in Weichang District, similar to that of present-day Zhaojue City, Sichuan Province in the Yangtze River Valley. When the palaeoclimatic parameters were compared with those of Middle Miocene Shanwang Basin, it would seem that the temperature and precipitation were a little higher in the Middle Miocene of eastern China. However, if the latitudinal temperature gradient at that time is considered, the median temperature values of GFY of Early Miocene and Shanwang of Mid-Miocene were similar.     

Dioscoreaceae fossils from the late Oligocene and early Miocene of Ethiopia

Dioscorea section Lasiophyton leaflets from the late Oligocene (27.23 Ma) and Tacca leaves from the early Miocene (21.73 Ma) of north‐western Ethiopia greatly expand the known fossil record of Dioscoreaceae and represent the earliest and only known records of the Afro‐Asian trifoliate, palmately veined yams (Dioscorea) and bat flowers (Tacca). Both fossils occur in volcaniclastic and clastic sediments associated with a high water table, and the palaeofloral assemblages are indicative of tropical moist forest formations. These fossils provide insight into the evolutionary history of the family in Africa during the mid‐Cenozoic and provide well‐dated taxa that can assist in phylogenetic analyses and evolutionary divergence studies for Dioscoreales and Dioscoreaceae. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 17–28.     

C4 photosynthesis evolved in warm climates but promoted migration to cooler ones

C₄ photosynthesis is considered an adaptation to warm climates, where its functional benefits are greatest and C₄ plants achieve their highest diversity and dominance. However, whether inherent physiological barriers impede the persistence of C₄ species in cool environments remains debated. Here, we use large grass phylogenetic and geographical distribution data sets to test whether (1) temperature influences the rate of C₄ origins, (2) photosynthetic types affect the rate of migration among climatic zones, and (3) C₄ evolution changes the breadth of the temperature niche. Our analyses show that C₄ photosynthesis in grasses originated in tropical climates, and that C₃ grasses were more likely to colonise cold climates. However, migration rates among tropical and temperate climates were higher in C₄ grasses. Therefore, while the origins of C₄ photosynthesis were concentrated in tropical climates, its physiological benefits across a broad temperature range expanded the niche into warmer climates and enabled diversification into cooler environments.     

Millennial-scale vegetation dynamics in an estuary at the onset of the Miocene Climate Optimum

Pollen analyses have been proven to possess the possibility to decipher rapid vegetational and climate shifts in Neogene sedimentary records. Herein, a c. 21-kyr-long transgression-regression cycle from the Lower Austrian locality Stetten is analysed in detail to evaluate climatic benchmarks for the early phase of the Middle Miocene Climate Optimum and to estimate the pace of environmental change.Based on the Coexistence Approach, a very clear signal of seasonality can be reconstructed. A warm and wet summer season with c. 204-236 mm precipitation during the wettest month was opposed by a rather dry winter season with precipitation of c. 9-24 mm during the driest month. The mean annual temperature ranged between 15.7 and 20.8 °C, with about 9.6-13.3 °C during the cold season and 24.7-27.9 °C during the warmest month. In contrast, today's climate of this area, with an annual temperature of 9.8 °C and 660 mm rainfall, is characterized by the winter season (mean temperature: -1.4 °C, mean precipitation: 39 mm) and a summer mean temperature of 19.9 °C (mean precipitation: 84 mm).Different modes of environmental shifts shaped the composition of the vegetation. Within few millennia, marshes and salt marshes with abundant Cyperaceae rapidly graded into Taxodiaceae swamps. This quick but gradual process was interrupted by swift marine ingressions which took place on a decadal to centennial scale. The transgression is accompanied by blooms of dinoflagellates and of the green alga Prasinophyta and an increase in Abies and Picea. Afterwards, the retreat of the sea and the progradation of estuarine and wetland settings were a gradual progress again.Despite a clear sedimentological cyclicity, which is related to the 21-kyr precessional forcing, the climate data show little variation. This missing pattern might be due to the buffering of the precessional-related climate signal by the subtropical vegetation. Another explanation could be the method-inherent broad range of climate-parameter estimates that could cover small scale climatic changes.     

Warm and humid Trans-Himalaya during the late Miocene: plant fossil evidence

Neogene fossil records from the Indus Basin sedimentary rocks (IBSR), deposited in the Indus Tsangpo Suture Zone (ITSZ), are very rare, but are important to understand the history of plant diversity and paleoclimate in the Himalaya. We report fossil wood ascribed to Ebenoxylon siwalicus Prakash from late Miocene sediments of the Karit Formation belonging to ITSZ. The anatomical details of the fossil wood, such as small to medium-sized vessels occluded with tyloses, scanty paratracheal to diffuse-in-aggregate axial parenchyma, 1–3 seriate homo to heterocellular rays, bordered intervessel pits with lenticular apertures and simple perforations, suggest its close affinity with Diospyros Linnaeus of the family Ebenaceae. Further anatomical details suggest a close resemblance with extant D. ehretioides Don and D. macrophylla Blume. The present fossil, along with previously known fossil records of Lagerstroemia (Lythraceae) and palms, indicate that the Trans-Himalaya was warm and humid during the late Miocene, quite different from the modern cool and dry climate in the study area.     

Miocene to Pliocene vegetation reconstruction and climate estimates in the Iberian Peninsula from pollen data

Pollen analysis of Miocene and Pliocene sediments from the Iberian Peninsula shows a progressive reduction in plant diversity through time caused by the disappearance of thermophilous and high-water requirement plants. In addition, an increase in warm-temperate (mesothermic), seasonal-adapted “Mediterranean” taxa, high-elevation conifers and herbs (mainly Artemisia) occurred during the Middle and Late Miocene and Pliocene. This has mainly been interpreted as a response of the vegetation to global and regional processes, including climate cooling related to the development of the East Antarctic Ice Sheet and then the onset of the Arctic Ice Sheet, uplift of regional mountains related to the Alpine uplift and the progressive movement of Eurasia towards northern latitudes as a result of the northwards subduction of Africa. The development of steppe-like vegetation in southern Iberia is ancient and probably started during the Oligocene. The onset of a contrasted seasonality in temperature during the Mid-Pliocene superimposed on the pre-existing seasonality in precipitation, the annual length of which increased southward. The Mediterranean climatic rhythm (summer drought) began about 3.4 Ma and caused the individualization of modern Mediterranean ecosystems. Quaternary-type Mediterranean climatic fluctuations started at 2.6 Ma (Gelasian) resulting in repeated steppe vs. forest alternations. A latitudinal climatic gradient between the southern and the northern parts of the Iberian Peninsula existed since the Middle Miocene.     

Fire does not alter vegetation in infertile prairie

The paradigm in prairie ecology is that fire is one of the key factors determining vegetation composition. Fire can impact grassland ecosystems in various ways, including changing plant species composition and inducing nitrogen loss. I found that 17 years of different burning frequencies in infertile grassland had only a minor impact on the vegetation composition and diversity. The only major impact from increasing the frequency of fires was a decrease of Poa pratensis abundance. However, other plant species did not respond to the change in Poa abundance. This result contrasts with previous studies in savannas and more productive grasslands, where the balance between trees, grasses, and the elimination of the litter layer can result in large vegetation changes. However, in this system primary productivity was low, litter did not accumulate and no major vegetation shifts occurred. Thus, the long-term vegetation impacts of burning in an infertile, low-productivity prairie were minimal.     

Timing of C4 grass expansion across sub-Saharan Africa

The emergence of C(4) grass biomes is believed to have first taken place in the upper Miocene, when a series of events modified global climate with long-lasting impacts on continental biotas. Changes included major shifts in floral composition-characterized in Africa by shrinking of forests and emergence of C(4) grasses and more open landscapes-followed by large-scale evolutionary shifts in faunal communities. The timing of the emergence of C(4) grasses, and the subsequent global expansion of C(4) grass-dominated biomes, however, is disputed, leading to contrasting views of the patterns of environmental changes and their links to faunal shifts, including those of early hominins. Here we evaluate the existing isotopic evidence available for central, eastern, and southern Africa, and review interpretations in light of these data. Pedogenic and biomineral carbonate delta(13)C data suggest that clear evidence for C(4) biomass in low latitudes exists only from 7-8 Ma. This likely postdates the emergence of C(4) plants, whose physiology is adapted to low atmospheric carbon dioxide concentrations. Biomes with C(4) grasses appeared later in mid-latitude sites. Moreover, C(4) grasses apparently remained a relatively minor component of most environments until the late Pliocene and early Pleistocene. Hence establishment of C(4) grasses, even as minor components of African biomes, precedes the very earliest evidence for bipedalism by two million years, and the more abundant and secure evidence by some three to four million years. This may suggest a protracted process of hominin adaptation to these emerging, more open landscapes.     

Fire severity affects vegetation and seed bank in a wetland

Questions: How does the severity of prescribed fires affect vegetation and seed bank in a wetland? Location: A fire‐prone reed swamp in northern Japan (250 ha, 40°49′N, 141°22′E, <10 m a.s.l.). Methods: Vegetation, biomass and seed bank were monitored for the 2 yr after annual prescribed fires were discontinued. Plant communities were placed into three categories based on fire severity: high (H) – fire consumed litter completely; moderate (M) – fire removed standing litter but left wet fallen litter; and low (L) – fire incompletely removed standing litter and did not remove fallen litter. Soil samples were collected in autumn 2007 and early summer 2008, and germinable seed bank was investigated by greenhouse trials. Results: High fire severity increased diversity in the next growing season by the establishment of short herbs in the standing vegetation. The biomass of forbs and grasses was greater in H where Phragmites australis biomass was reduced. The density of seed bank was >30 000 seeds m^?2 throughout all the treatments. Perennial plants were dominant in the vegetation, while annuals, biennials and rushes were dominant in the seed bank. Small seeds were more abundant in the soil than in the litter. Qualitative and quantitative similarities between seed bank and the vegetation were low, and tended to be higher in H. Conclusions: Fire contributed to the development of diverse standing vegetation via the positive effects on seed bank dynamics, and can be considered a tool to maintain species‐rich marshes.