Toward a rule-based biome model

EXTRACT (SEE PDF FOR FULL ABSTRACT): Current projections of the response of the biosphere to global climatic change indicate as much as 50 to 90% spatial displacement of extratropical biomes. The mechanism of spatial shift could be dominated either by competitive displacement of northern biomes by southern biomes or by drought-induced dieback of areas susceptible to change. The current suite of global biosphere models cannot distinguish between these two processes, hence the need for a mechanistically based biome model. The first steps have been taken toward development of a rule-based, mechanistic model of regional biomes at a continental scale. ... The model is in an early stage of development and will require several enhancements, including: explicit simulation of potential evapotranspiration, extension to boreal and tropical biomes, a shift from steady-state to transient dynamics, and validation on other continents.

Pollen-based biome reconstructions for Latin America at 0, 6000 and 18000 radiocarbon years

The biomisation method is used to reconstruct Latin American vegetation at 6000±500 and 18 000±1000 radiocarbon years before present (14C yr BP) from pollen data. Tests using modern pollen data from 381 samples derived from 287 locations broadly reproduce potential natural vegetation. The strong temperature gradient associated with the Andes is recorded by a transition from high altitude cool grass/shrubland and cool mixed forest to mid-altitude cool temperate rain forest, to tropical dry, seasonal and rain forest at low altitudes. Reconstructed biomes from a number of sites do not match the potential vegetation due to local factors such as human impact, methodological artefacts and mechanisms of pollen representivity of the parent vegetation.

Palaeovegetation in China during the late Quaternary: Biome reconstructions based on a global scheme of plant functional types

Two previous reconstructions of palaeovegetation across the whole of China were performed using a simple classification of plant functional types (PFTs). Now a more explicit, global PFT classification scheme has been developed, and a substantial number of additional pollen records have become available. Here we apply the global scheme of PFTs to a comprehensive set of pollen records available from China to test the applicability of the global scheme of PFTs in China, and to obtain a well-founded reconstruction of changing palaeovegetation patterns. A total of 806 pollen surface samples, 188 mid-Holocene (MH, 6000 14C yr BP) and 50 last glacial maximum (LGM, 18,000 14C yr BP) pollen records were used to reconstruct vegetation patterns in China, based on a new global classification system of PFTs and a standard numerical technique for biome assignment (biomization). The biome reconstruction based on pollen surface samples showed convincing agreement with present potential natural vegetation. Coherent patterns of change in biome distribution between MH, LGM and present are observed. In the MH, cold and cool-temperate evergreen needleleaf forests and mixed forests, temperate deciduous broadleaf forest, and warm-temperate evergreen broadleaf and mixed forest in eastern China were shifted northward by 200–500 km. Cold-deciduous forest in northeastern China was replaced by cold evergreen needleleaf forest while in central northern China, cold-deciduous forest was present at some sites now occupied by temperate grassland and desert. The forest–grassland boundary was 200–300 km west of its present position. Temperate xerophytic shrubland, temperate grassland and desert covered a large area on the Tibetan Plateau, but the area of tundra was reduced. Treeline was 300–500 m higher than present in Tibet. These changes imply generally warmer winters, longer growing seasons and more precipitation during the MH. Westward shifts of the forest–shrubland–grassland and grassland–desert boundaries imply greater moisture availability in the MH, consistent with a stronger summer monsoon. During the LGM, in contrast, cold-deciduous forest, cool-temperate evergreen needleleaf forest, cool mixed forests, warm-temperate evergreen broadleaf and mixed forest in eastern China were displaced to the south by 300–1000 km, while temperate deciduous broadleaf forest, pure warm-temperate evergreen forest, tropical semi-evergreen and evergreen broadleaf forests were restricted or absent from the mainland of southern China, implying colder winters than present. Strong shifts of temperate xerophytic shrubland, temperate grassland and desert to the south and east in northern and western China and on the Tibetan Plateau imply drier conditions than present.

Pollen-based reconstructions of biome distributions for Australia, Southeast Asia and the Pacific (SEAPAC region) at 0,6000 and 18,000 14C yr B.P.

Aim  This paper documents reconstructions of the vegetation patterns in Australia, Southeast Asia and the Pacific (SEAPAC region) in the mid-Holocene and at the last glacial maximum (LGM). Methods  Vegetation patterns were reconstructed from pollen data using an objective biomization scheme based on plant functional types. The biomization scheme was first tested using 535 modern pollen samples from 377 sites, and then applied unchanged to fossil pollen samples dating to 6000 ± 500 or 18,000 ± 1000 14C yr bp. Results  1. Tests using surface pollen sample sites showed that the biomization scheme is capable of reproducing the modern broad-scale patterns of vegetation distribution. The north–south gradient in temperature, reflected in transitions from cool evergreen needleleaf forest in the extreme south through temperate rain forest or wet sclerophyll forest (WSFW) and into tropical forests, is well reconstructed. The transitions from xerophytic through sclerophyll woodlands and open forests to closed-canopy forests, which reflect the gradient in plant available moisture from the continental interior towards the coast, are reconstructed with less geographical precision but nevertheless the broad-scale pattern emerges. 2. Differences between the modern and mid-Holocene vegetation patterns in mainland Australia are comparatively small and reflect changes in moisture availability rather than temperature. In south-eastern Australia some sites show a shift towards more moisture-stressed vegetation in the mid-Holocene with xerophytic woods/scrub and temperate sclerophyll woodland and shrubland at sites characterized today by WSFW or warm-temperate rain forest (WTRF). However, sites in the Snowy Mountains, on the Southern Tablelands and east of the Great Dividing Range have more moisture-demanding vegetation in the mid-Holocene than today. South-western Australia was slightly drier than today. The single site in north-western Australia also shows conditions drier than today in the mid-Holocene. Changes in the tropics are also comparatively small, but the presence of WTRF and tropical deciduous broadleaf forest and woodland in the mid-Holocene, in sites occupied today by cool-temperate rain forest, indicate warmer conditions. 3. Expansion of xerophytic vegetation in the south and tropical deciduous broadleaf forest and woodland in the north indicate drier conditions across mainland Australia at the LGM. None of these changes are informative about the degree of cooling. However the evidence from the tropics, showing lowering of the treeline and forest belts, indicates that conditions were between 1 and 9 °C (depending on elevation) colder. The encroachment of tropical deciduous broadleaf forest and woodland into lowland evergreen broadleaf forest implies greater aridity. Main conclusions  This study provides the first continental-scale reconstruction of mid-Holocene and LGM vegetation patterns from Australia, Southeast Asia and the Pacific (SEAPAC region) using an objective biomization scheme. These data will provide a benchmark for evaluation of palaeoclimate simulations within the framework of the Palaeoclimate Modelling Intercomparison Project.

Climate and CO2 controls on global vegetation distribution at the last glacial maximum: analysis based on palaeovegetation data, biome modelling and palaeoclimate simulations.

The global vegetation response to climate and atmospheric CO2 changes between the last glacial maximum and recent times is examined using an equilibrium vegetation model (BIOME4), driven by output from 17 climate simulations from the Palaeoclimate Modelling Intercomparison Project. Features common to all of the simulations include expansion of treeless vegetation in high northern latitudes; southward displacement and fragmentation of boreal and temperate forests; and expansion of drought-tolerant biomes in the tropics. These features are broadly consistent with pollen-based reconstructions of vegetation distribution at the last glacial maximum. Glacial vegetation in high latitudes reflects cold and dry conditions due to the low CO2 concentration and the presence of large continental ice sheets. The extent of drought-tolerant vegetation in tropical and subtropical latitudes reflects a generally drier low-latitude climate. Comparisons of the observations with BIOME4 simulations, with and without consideration of the direct physiological effect of CO2 concentration on C3 photosynthesis, suggest an important additional role of low CO2 concentration in restricting the extent of forests, especially in the tropics. Global forest cover was overestimated by all models when climate change alone was used to drive BIOME4, and estimated more accurately when physiological effects of CO2 concentration were included. This result suggests that both CO2 effects and climate effects were important in determining glacial-interglacial changes in vegetation. More realistic simulations of glacial vegetation and climate will need to take into account the feedback effects of these structural and physiological changes on the climate.

Pollen-based biome reconstructions for China at 0 and 6000 years

Biomization provides an objective and robust method of assigning pollen spectra to biomes so that pollen data can be mapped and compared directly with the output of biomgeographic models. We have tested the applicability of this procedure, originally developed for Europe, to assign modern surface samples from China to biomes. The procedure successfully delineated the major vegetation types of China. When the same procedure was applied to fossil pollen samples for 6000 years ago, the reconstructions showed systematic differences from present, consistent with previous interpretations of vegetation changes since the mid-Holocene. In eastern China, the forest zones were systematically shifted northwards, such that cool mixed forests displaced taiga in northeastern China, while broad-leaved evergreen forest extended c. 300 km and temperate deciduous forestc. 500–600 km beyond their present northern limits. In northwestern China, the area of desert and steppe vegetation was reduced compared to present. On the Tibetan Plateau, forest vegetation extended to higher elevations than today and the area of tundra was reduced. These shifts in biome distributions imply significant changes in climate since 6000 years ago that can be interpreted qualitatively as a response to orbital forcing and its secondary effects on the Asian monsoon.

Climate and biome simulations for the past 21,000 years

This paper reports on a set of paleoclimate simulations for 21, 16, 14, 11 and 6 ka (thousands of years ago) carried out with the Community Climate Model, Version 1 (CCM1) of the National Center for Atmospheric Research (NCAR). This climate model uses four interactive components that were not available in our previous simulations with the NCAR CCM0 (COHMAP, 1988Science, 241, 1043–1052; Wright et al., 1993Global Climate Since the Last Glocial Maximum, University of Minnesota Press, MN): soil moisture, snow hydrology, sea-ice, and mixed-layer ocean temperature. The new simulations also use new estimates of ice sheet height and size from ( Peltier 1994, Science, 265, 195–201), and synchronize the astronomically dated orbital forcing with the ice sheet and atmospheric CO2 levels corrected from radiocarbon years to calendar years. The CCM1 simulations agree with the previous simulations in their most general characteristics. The 21 ka climate is cold and dry, in response to the presence of the ice sheets and lowered CO2 levels. The period 14–6 ka has strengthened northern summer monsoons and warm mid-latitude continental interiors in response to orbital changes. Regional differences between the CCM1 and CCM0 simulations can be traced to the effects of either the new interactive model components or the new boundary conditions. CCM1 simulates climate processes more realistically, but has additional degrees of freedom that can allow the model to ‘drift’ toward less realistic solutions in some instances. The CCM1 simulations are expressed in terms of equilibrium vegetation using BIOME 1, and indicate large shifts in biomes. Northern tundra and forest biomes are displaced southward at glacial maximum and subtropical deserts contract in the mid-Holocene when monsoons strengthen. These vegetation changes could, if simulated interactively, introduce additional climate feedbacks. The total area of vegetated land remains nearly constant through time because the exposure of continental shelves with lowered sea level largely compensates for the land covered by the expanded ice sheets.

Comparing projections of future changes in runoff from hydrological and biome models in ISI-MIP

Future changes in runoff can have important implications for water resources and flooding. In this study, runoff projections from ISI-MIP (Inter-sectoral Impact Model Inter-comparison Project) simulations forced with HadGEM2-ES bias-corrected climate data under the Representative Concentration Pathway 8.5 have been analysed for differences between impact models. Projections of change from a baseline period (1981-2010) to the future (2070-2099) from 12 impacts models which contributed to the hydrological and biomes sectors of ISI-MIP were studied. The biome models differed from the hydrological models by the inclusion of CO2 impacts and most also included a dynamic vegetation distribution. The biome and hydrological models agreed on the sign of runoff change for most regions of the world. However, in West Africa, the hydrological models projected drying, and the biome models a moistening. The biome models tended to produce larger increases and smaller decreases in regionally averaged runoff than the hydrological models, although there is large inter-model spread. The timing of runoff change was similar, but there were differences in magnitude, particularly at peak runoff. The impact of vegetation distribution change was much smaller than the projected change over time, while elevated CO2 had an effect as large as the magnitude of change over time projected by some models in some regions. The effect of CO2 on runoff was not consistent across the models, with two models showing increases and two decreases. There was also more spread in projections from the runs with elevated CO2 than with constant CO2. The biome models which gave increased runoff from elevated CO2 were also those which differed most from the hydrological models. Spatially, regions with most difference between model types tended to be projected to have most effect from elevated CO2, and seasonal differences were also similar, so elevated CO2 can partly explain the differences between hydrological and biome model runoff change projections. Therefore, this shows that a range of impact models should be considered to give the full range of uncertainty in impacts studies.

Patterns of water and heat flux across a biome gradient from tropical forest to savanna in Brazil

We investigated the seasonal patterns of water vapor and sensible heat flux along a tropical biome gradient from forest to savanna. We analyzed data from a network of flux towers in Brazil that were operated within the Large-Scale Biosphere-Atmosphere Experiment in Amazonia (LBA). These tower sites included tropical humid and semideciduous forest, transitional forest, floodplain (with physiognomies of cerrado), and cerrado sensu stricto. The mean annual sensible heat flux at all sites ranged from 20 to 38 Wm(-2), and was generally reduced in the wet season and increased in the late dry season, coincident with seasonal variations of net radiation and soil moisture. The sites were easily divisible into two functional groups based on the seasonality of evaporation: tropical forest and savanna. At sites with an annual precipitation above 1900 mm and a dry season length less than 4 months (Manaus, Santarem and Rondonia), evaporation rates increased in the dry season, coincident with increased radiation. Evaporation rates were as high as 4.0 mm d(-1) in these evergreen or semidecidous forests. In contrast, ecosystems with precipitation less than 1700 mm and a longer dry season (Mato Grosso, Tocantins and Sao Paulo) showed clear evidence of reduced evaporation in the dry season. Evaporation rates were as low as 2.5 mm d(-1) in the transitional forests and 1 mm d(-1) in the cerrado. The controls on evapotranspiration seasonality changed along the biome gradient, with evaporative demand (especially net radiation) playing a more important role in the wetter forests, and soil moisture playing a more important role in the drier savannah sites.

Birth of a biome : insights into the assembly and maintenance of the Australian arid zone biota

The integration of phylogenetics, phylogeography and palaeoenvironmental studies is providing major insights into the historical forces that have shaped the Earth’s biomes. Yet our present view is biased towards arctic and temperate/tropical forest regions, with very little focus on the extensive arid regions of the planet. The Australian arid zone is one of the largest desert landform systems in the world, with a unique, diverse and relatively well-studied biota. With foci on palaeoenvironmental and molecular data, we here review what is known about the assembly and maintenance of this biome in the context of its physical history, and in comparison with other mesic biomes. Aridification of Australia began in the Mid-Miocene, around 15 million years, but fully arid landforms in central Australia appeared much later, around 1–4 million years. Dated molecular phylogenies of diverse taxa show the deepest divergences of arid-adapted taxa from the Mid-Miocene, consistent with the onset of desiccation. There is evidence of arid-adapted taxa evolving from mesicadapted ancestors, and also of speciation within the arid zone. There is no evidence for an increase in speciation rate during the Pleistocene, and most arid-zone species lineages date to the Pliocene or earlier. The last 0.8 million years have seen major fluctuations of the arid zone, with large areas covered by mobile sand dunes during glacial maxima. Some large, vagile taxa show patterns of recent expansion and migration throughout the arid zone, in parallel with the ice sheet-imposed range shifts in Northern Hemisphere taxa. Yet other taxa show high lineage diversity and strong phylogeographical structure, indicating persistence in multiple localised refugia over several glacial maxima. Similar to the Northern Hemisphere, Pleistocene range shifts have produced suture zones, creating the opportunity for diversification and speciation through hybridisation, polyploidy and parthenogenesis. This review highlights the opportunities that development of arid conditions provides for rapid and diverse evolutionary radiations, and re-enforces the emerging view that Pleistocene environmental change can have diverse impacts on genetic structure and diversity in different biomes. There is a clear need for more detailed and targeted phylogeographical studies of Australia’s arid biota and we suggest a framework and a set of a priori hypotheses by which to proceed.

Habitat use and home range of brown-nosed coati, Nasua nasua (Carnivora: Procyonidae) in the Brazilian Cerrado biome

The brown-nosed coati (Nasua nasua) is a carnivorous species found in all the Brazilian biomes, some of which are endangered areas. The aim of this work was to determine the habitat use and selection, home range and core area of N. nasua in the Cerrado biome, central region of Tocantins, Brazil. The study was carried out in an area of approximately 20 000ha from May 2000 to July 2002. A total of seven box traps were placed in the area for 13 months, three of 11 captured animals were followed and monitored by radio-tracking during 13 months. The monitoring was conducted once a day, three times a week using a car and walking through the study area (radio-tracking and visual contact). The results demonstrate that these three males used more frequently the gallery forest formation, followed by cerrado and wetlands. The use of gallery forest by these animals indicated an habitat selection (Proportion test, z=12.98, p< 0.01). Besides, adult males used the gallery forest more frequently (Fisher's exact test, p<0.01) and wetlands less frequently (Fisher's exact test, p<0.01) than juvenile males, without significant differences between animal ages for cerrado percentage of habitat use. Besides, results also showed a gallery forest selection by adult (Proportion test z= 13.62, p<0.01) and juvenile (Proportion test z=2.68, p<0.01) males, and a wetland selection by the juvenile male (Proportion test z=3.90, p<0.01). The home ranges varied from 2.20 to 7.55km2 for the Minimum Convex Polygon 100% (MCP 100%) and from 4.38 to 13.32km2 for the Harmonic Mean 95% (HM 95%). The smallest home range overlap occurred between the adult males (Nm1 and Nm3), and the greatest between the juvenile Njm2 and the adult Nm1. The average of the core area (HM 75%) for the three monitored animals represented 21.29% of the home range calculated with HM 95%. No overlap between core areas was observed for adult males, but, it was an overlap between the core area of the juvenile male and its band with that of the two adult males. The present study provides new data on core area size and frequency habitat use by adult and juvenile males of N. nasua in the Brazilian Cerrado, that may support conservation efforts. Rev. Biol. Trop. 58 (3): 1069-1077. Epub 2010 September 01.

HELMINTH FAUNA of TWO GECKO LIZARDS, HEMIDACTYLUS AGRIUS and LYGODACTYLUS KLUGEI (GEKKONIDADE), FROM CAATINGA BIOME, NORTHEASTERN BRAZIL.

This study presented data on helminth fauna of two gecko lizards, Hemidactylus agrius and Lygodactylus klugei, from Caatinga biome in northeastern Brazil. It was found four helminth species parasitizing H. agrius, cistacanth of Centrorhynchidae (Acanthocephala) and the nematodes Physalopteridae (larvae), Parapharyngodon alvarengai (Pharyngodonidae) and Skrjabinelazia sp. (Seuratidade). The host Lygodactylus klugei presented two helminth species, one individual of Mesocoelium monas (Trematoda: Mesocoeliidae) in the small intestine and one encysted larvae of Physalopteridae (Nematoda: Physalopteridae) attached at stomach wall. The lizard species showed a low prevalence and low richness of helminths. Moreover, H. agrius presented a low intensity of infection. The foraging mode, arboreal habit and a restricted composition of diet could favoring the low prevalence, low infection rates and low richness of helminths found in these geckonid host species.

Nematodes of Lizards (Reptilia: Squamata) from Caatinga Biome, Northeastern Brazil

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)