Biodiversity from a historical geology perspective: a case study from Marajó Island, lower Amazon

Assessing patterns of abundance and distribution of Amazonian species is still an overwhelming task that requires integration of multiple disciplines. This work is based on background information gathered from previous reconstructions of the geological history of the lower Amazon drainage basin, in order to analyse biodiversity patterns within the context of landscape transformation. A highly dynamic geological scenario is depicted for this area during the Plio‐Pleistocene and Holocene, which consisted of a large palaeovalley formed as a response of tectonic reactivation. This palaeovalley was filled with sediments transported by a north/northwest orientated palaeo‐Tocantins River. The palaeodrainage became abandoned as the main river course was deviated to the northeast, initiating the separation of Marajó Island from mainland. Geology had a direct impact on the modern physiognomy, with open vegetation dominating in areas with Holocene sedimentation, while closed forests prevailing in older Quaternary and, probably also, Pliocene terrains. Data from fossil and modern mammalian groups indicate the connection of Marajó Island to the mainland during the Last Glacial Maximum, when open vegetation seems to have dominated. Tectonic subsidence was responsible for the maintenance of this vegetation pattern on the eastern side of the Marajó Island, keeping it as a habitat favourable for savanna adapted faunal elements. Based on this kind of information, this work attempts to highlight the importance of integrating studies combining geological and biological events as the key to understand biodiversity patterns in Amazonia. It is expected to open new lines of research dealing with the comprehension of ecology, species and genetic diversity, biogeography, evolutionary scenarios, and speciation mechanisms.     

Stable forest–savanna mosaic in north‐western Tanzania: local‐scale evidence from δ13C signatures and 14C ages of soil fractions

Aim The spatio‐temporal dynamics of dry evergreen forest patches in the savanna biome of the Kagera region (north‐western Tanzania) are largely unknown owing to a lack of pollen and macrofossil evidence. Our aims were to reconstruct local‐scale shifts of the forest–savanna boundary in order to determine whether the forests have been expanding or retreating on a centennial and millennial time‐scale. Location The Kagera region of north‐western Tanzania, East Africa. Methods The vegetation reconstruction was based on analysing δ¹³C signatures in soils along a transect spanning both C₄ open savanna and C₃ forest vegetation. Furthermore, we fractionated soil organic matter (SOM) according to density and chemical stability to analyse δ¹³C values of soil fractions with distinct radiocarbon ages. Results We found sharp changes in δ¹³C signatures in bulk SOM from the forest to the savanna, within a few metres along the transect. The forest soil profiles carried a persistent C₃‐dominated signature. Radiocarbon dating of the oldest, most recalcitrant forest soil fraction yielded a mean age of 5500 cal. yr bp , demonstrating that the forest has existed since at least the mid‐Holocene. The savanna sites showed a typical C₄ isotopic signature in SOM of topsoils, but subsoils and more recalcitrant SOM fractions also contained signals of C₃ plants. The dense soil fraction (ρ > 1.6 g cm^?3) carrying a pure C₄ label had a mean age of c. 1200 cal. yr bp , indicating the minimum duration of the dominance of grass vegetation on the savanna site. At the forest edge, the older C₄ grass signature of SOM has steadily been replaced by the more negative δ¹³C fingerprint of the forest trees. As this replacement has occurred mainly in the 10‐m‐wide forest–savanna ecotone over the last c. 1200 years, the forest expansion must be very slow and is very likely less than 15 m century^?1. Main conclusions Our results suggest that forest patches in the Kagera savanna landscape are very stable vegetation formations which have persisted for millennia. During the last millennium, they have been expanding very slowly into the surrounding savanna at a rate of less than 15 m century^?1.     

Patterns of plant phenology in Amazonian seasonally flooded and unflooded forests

Few studies have successfully monitored community‐wide phenological patterns in seasonally flooded Amazonian várzea forests, where a prolonged annual flood pulse arguably generates the greatest degree of seasonality of any low‐latitude ecosystem on Earth. We monitored the vegetative and reproductive plant phenology of várzea (VZ) floodplain and adjacent terra firme (TF) forests within two contiguous protected areas in western Brazilian Amazonia, using three complementary methods: monthly canopy observations of 1056 individuals (TF: 556, VZ: 500), twice monthly collections from 0.5‐m² litterfall traps within two 100‐ha plots (1 TF, 1 VZ; 96 traps per plot), and monthly ground surveys of residual fruit‐fall along transect‐grids within each 100‐ha plot (12 km per plot). Surveys encompassed the entire annual flood cycle and employed a floating trap design to cope with fluctuating water levels. Phenology patterns were generally similar in both forest types. Leaffall peaked during the aquatic phase in várzea forest and the dry season in terra firme. Flowering typically followed leaffall and leaf flush, extending into the onset of the terrestrial phase and rainy season in várzea and terra firme, respectively. Abiotic seed dispersal modes were relatively more prevalent in várzea than terra firme; the main contrast in fruiting seasonality was more likely a result of differences in community composition and relative abundance of seed dispersal modes than differences within individual genera. We emphasize the difficulty in distinguishing the role of the flood pulse from other seasonal environmental variables without multiannual data or spatially replicated studies across the spectrum of Amazonian forest types.     

Phyllostomid Bat Assemblage Structure in Amazonian Flooded and Unflooded Forests

In Amazonia, the assemblages of several taxa differ significantly between upland terra firme and white‐water flooded várzea forests, but little is known about the diversity and distribution of bats in these two forest types. We compare the spatio‐temporal patterns of bat assemblage composition and structure in adjacent terra firme and várzea forests in the lower Purus River region of central Brazilian Amazonia. Bats were sampled using mist nets at five sites in each forest type during 40 nights (2400 net‐hours). We captured 1069 bats representing 42 species and Phyllostomidae bats comprised 99.3 percent of all captures. The bat assemblages in várzea and terra firme forests were significantly different, mainly due to a marked dissimilarity in species composition and in the number of captures during high‐water season. In addition, bat assemblages within forest types differed significantly between seasons for both terra firme and várzea. Frugivores dominated the bat assemblages in both forest types. Overall guild structure did not change between várzea and terra firme or between seasons, but frugivore and animalivore abundance increased significantly in várzea forest during the inundation. The difference in assemblage structure observed in the high‐water season is probably caused by the annual várzea flooding, which provides an effective barrier to the persistence of many understory bats. We also hypothesize that some bat species may undertake seasonal movements between forest types in response to fruit abundance, and our results further underline the importance of floodplain habitats for the conservation of species in the Amazon.     

Seasonal flooding decreases fruit-feeding butterfly species dominance and increases spatial turnover in floodplain forests of central Amazonia

The seasonal flood pulse in Amazonia can be considered a primary driver of community structure in floodplain environments. Although this natural periodic disturbance is part of the landscape dynamics, the seasonal inundation presents a considerable challenge to organisms that inhabit floodplain forests. The present study investigated the effect of seasonal flooding on fruit-feeding butterfly assemblages in different forest types and strata in central Amazonia. We sampled fruit-feeding butterflies in the canopy and the understory using baited traps in adjacent upland (unflooded forests—terra firme), white and blackwater floodplain forests (várzea and igapó, respectively) during the low- and high-water seasons. Butterfly abundance decreased in the high-water season, especially of dominant species in várzea, but the number of species was similar between seasons in the three forest types. Species composition differed between strata in all forest types. However, the flood pulse only affected butterfly assemblages in várzea forest. The β-diversity components also differed only in várzea. Species replacement (turnover) dominated the spatial β-diversity in igapó and terra firme in both seasons and várzea in the high-water season. Nonetheless, nestedness was relatively higher in várzea forests during the low-water season, mainly due to the effect of dominant species. These results emphasize the importance of seasonal flooding to structure butterfly assemblages in floodplain forests and reveal the idiosyncrasy of butterfly community responses to flooding in different forest types. Our results also suggest that any major and rapid changes to the hydrological regime could severely affect floodplain communities adapted to this natural seasonal hydrological cycle, threatening the existence of these unique environments.     

Vertebrate responses to fruit production in Amazonian flooded and unflooded forests

We document patterns of fruit and vertebrate abundance within an extensive, virtually undisturbed mosaic of seasonally flooded (várzea and igapó) and unflooded (terra firme) forests of central Amazonia. Using phenological surveys and a standardised series of line-transect censuses we investigate the spatial and temporal patterns of immature and mature fruit availability and how this may affect patterns of habitat use by vertebrates in the landscape. All habitats showed marked peaks in fruiting activity, and vertebrate detection rates varied over time for most species both within and between forest types. Many arboreal and terrestrial vertebrates used both types of flooded forest on a seasonal basis, and fluctuations in the abundance of terrestrial species in várzea forest were correlated with fruit availability. Similarly, the abundance of arboreal seed predators such as buffy saki monkeys (Pithecia albicans) and macaws (Ara spp.) were closely linked with immature fruit availability in terra firme forest. We conclude that highly heterogeneous landscapes consisting of terra firme, várzea and igapó forest appear to play an important role in the dynamics of many vertebrate species in lowland Amazonia, but the extent to which different forest types are used is highly variable in both space and time.     

Functional differences between woodland savannas and seasonally dry forests from south‐eastern Brazil: Evidence from 15N natural abundance studies

Nitrogen availability and N‐cycling dynamics across ecosystems play a critical role in plant functioning and species distribution. Measurements of ¹⁵N natural abundance provides a way to assess ecosystem N dynamics, and the range of nitrogen stable isotope values (δ¹⁵N) for plants in an ecosystem can indicate divergent strategies for N uptake. We tested the hypotheses that the N‐rich seasonally dry forest would have higher soil and leaf δ¹⁵N and a smaller range of leaf δ¹⁵N values compared to the N‐poor cerradão (savanna woodland). We measured N concentration and δ¹⁵N in two soil depths and leaves of 27 woody species in cerradão and 26 in seasonally dry forest. As expected, total soil N concentration decreased while soil δ¹⁵N value increased with soil depth. Regardless of soil depth, seasonally dry forest soils had higher δ¹⁵N and total N concentration compared to cerradão soils. Foliar δ¹⁵N values varied from ?6.4‰ to 5.9‰ in cerradão and from ?2.3‰ to 8.4‰ in seasonally dry forest plants. Phylogenetically independent contrasts analysis and comparisons of δ¹⁵N mean values of the most abundant species and species co‐occurring in both sites confirmed the hypothesis of higher δ¹⁵N for seasonally dry forest in comparison to cerradão. These results corroborate the expectation of higher soil and leaf δ¹⁵N values in sites with higher soil N availability. However, except for the most abundant species, no across‐site leaf–soil (δ¹⁵N leaf –δ¹⁵N soil) differences (Δδ¹⁵N) were found suggesting that differences in leaf δ¹⁵N between cerradão and seasonally dry forest are driven by differences in soil δ¹⁵N. Variation of leaf δ¹⁵N was large in both sites and only slightly higher in cerradão, suggesting high diversity of N use strategies for both cerradão and seasonally dry forest communities.     

Tree species composition and diversity gradients in white-water forests across the Amazon Basin

Aim Attention has increasingly been focused on the floristic variation within forests of the Amazon Basin. Variations in species composition and diversity are poorly understood, especially in Amazonian floodplain forests. We investigated tree species composition, richness and α diversity in the Amazonian white‐water (várzea) forest, looking particularly at: (1) the flood‐level gradient, (2) the successional stage (stand age), and (3) the geographical location of the forests. Location Eastern Amazonia, central Amazonia, equatorial western Amazonia and the southern part of western Amazonia. Methods The data originate from 16 permanent várzea forest plots in the central and western Brazilian Amazon and in the northern Bolivian Amazon. In addition, revised species lists of 28 várzea forest inventories from across the Amazon Basin were used. Most important families and species were determined using importance values. Floristic similarity between plots was calculated to detect similarity variations between forest types and over geographical distances. To check for spatial diversity gradients, α diversity (Fisher) of the plots was correlated with stand age, longitudinal and latitudinal plot location, and flood‐level gradient. Results More than 900 flood‐tolerant tree species were recorded, which indicates that Amazonian várzea forests are the most species‐rich floodplain forests worldwide. The most important plant families recorded also dominate most Neotropical upland forests, and c. 31% of the tree species listed also occur in the uplands. Species distribution and diversity varied: (1) on the flood‐level gradient, with a distinct separation between low‐várzea forests and high‐várzea forests, (2) in relation to natural forest succession, with species‐poor forests in early stages of succession and species‐rich forests in later stages, and (3) as a function of geographical distance between sites, indicating an increasing α diversity from eastern to western Amazonia, and simultaneously from the southern part of western Amazonia to equatorial western Amazonia. Main conclusions The east‐to‐west gradient of increasing species diversity in várzea forests reflects the diversity patterns also described for Amazonian terra firme. Despite the fine‐scale geomorphological heterogeneity of the floodplains, and despite high disturbance of the different forest types by sedimentation and erosion, várzea forests are dominated by a high proportion of generalistic, widely distributed tree species. In contrast to high‐várzea forests, where floristic dissimilarity increases significantly with increasing distance between the sites, low‐várzea forests can exhibit high floristic similarity over large geographical distances. The high várzea may be an important transitional zone for lateral immigration of terra firme species to the floodplains, thus contributing to comparatively high species richness. However, long‐distance dispersal of many low‐várzea trees contributes to comparatively low species richness in highly flooded low várzea.     

Can stable carbon isotopes (δ13C) in soil carbon be used to describe the dynamics of Eucalyptus savanna–rainforest boundaries in the Australian monsoon tropics?

The history of isolated patches of monsoon rainforest within large tracts of Eucalyptus savanna is poorly understood because of the scarcity of reliable palaeoecological records in the Australian monsoon tropics. Elsewhere in the world, the ratio of the stable isotopes ¹³C to ¹²C (δ¹³C) in soil organic matter has shed light on the dynamics of rainforest–savanna boundaries because tropical grasses with the C4 photosynthetic pathway have a distinct δ¹³C signature (–17 to –9‰) compared with that of woody plants with the C3 photosynthetic pathway (–32 to –22‰). In order to determine the magnitude of the variation in δ¹³C, unreplicated soil profiles were sampled beneath different vegetation types on three boundaries between Eucalyptus savanna and rainforest that were both growing on Tertiary age laterite parent material. Replicated (n = 3) soil profiles, which were also derived from Tertiary age laterite, were sampled from beneath: (i) dense stands of African grasses within a frequently burnt Eucalyptus savanna; and within the same long unburnt Eucalyptus savanna, (ii) patches of African and natives grasses and (iii) clumps of Acacia trees. The strongly negative δ¹³C values of soil organic matter derived from the frequently burnt and long unburnt grassy understoreys in the Eucalyptus savannas showed that a considerable amount of the soil carbon was derived from C3 (woody) species despite the presence of a ground layer dominated by C4 grasses. However, a feature of these data was the considerable variability among the three ‘replicate’ profiles. The surface soil samples from beneath three clumps of Acacia trees in the unburnt Eucalyptus savanna had much less variable δ¹³C values and were similar to two of the three monsoon rainforests sampled. The pattern of δ¹³C values from unreplicated soil profiles from different vegetation types across three rainforest boundaries was also very variable and not always obviously related the known disturbance history of the extant vegetation. Given the considerable variability within and between vegetation types with contrasting disturbance histories, it is concluded that the use of carbon stable isotopes to advance understanding of the dynamics of rainforest and Eucalyptus savanna boundaries will require further development, such as determination of the ¹⁴C age and δ¹³C values of different soil carbon fractions.     

Soil seed banks in different environments: initial forest,mature forest,Pinus and Eucalyptus abandoned stands

Abstract

Different environments (initial forest, mature forest, Pinus and Eucalyptus stands) found in Seasonal Semideciduous Forest fragments affect the density of viable seeds, as well as the floristic similarity, diversity, and richness of tree species in soil seed banks. This hypothesis was tested in the current study. Soil seed bank samples were collected in the aforementioned environments during rainy and dry seasons, and taken to a shade house, where they remained under favorable seed germination conditions. Tree seedling emergence was measured, and sample species were identified every 15?days, for six months, in each sampling period. In total, 97 individuals m^?2 and 23 species emerged in all environments and periods. The highest density of viable seeds of tree species in the soil seed bank was found in the initial forest stretch, mature forest stretch and abandoned Eucalyptus stand. Only the Pinus stand seed bank in the dry season had different floristic and lower viable seed density than the mature forest seed bank. Thus, all environments, except the abandoned Pinus stand, can preserved Seasonal Semideciduous Forest fragments.     

Not all forests are expanding over central Brazilian savannas

Recently we reported on the expansion of riparian forests into savannas in central Brazil. To enlarge the scope of the earlier study we investigated whether upland deciduous and xeromorphic forests behaved similarly. We investigated past vegetation changes that occurred in forest/savanna transitions using carbon isotope ratios (δ¹³C) measured in the soil organic matter as a tracer. We analyzed the ¹⁴C activity where δ¹³C showed major shifts in vegetation. The role of soil chemical and physical attributes in defining vegetation distribution is discussed. Structural changes in vegetation were found to be associated with shifts in the isotope composition (δ¹³C) of soil organic matter. This was attributed to intrinsic differences in the biomass of trees and grasses and allowed for the determination of past shifts in vegetation by evaluating δ¹³C at different depths. The deciduous forest decreased in area approximately 980 years ago. Tree cover increased in the xeromorphic forest, but the border stayed stable through time. The deciduous forest and adjacent savanna have eutrophic soils while the xeromorphic forest and adjacent savanna have dystrophic soils. However, greater organic carbon, nitrogen and phosphorus concentrations are observed in the forests. We provide concrete evidence of deciduous forest retreat unlike the stability observed in the xeromorphic forest/savanna boundary. These results contrast with the expansion of riparian forests recently reported in the same region.     

Notes on the geographic distribution of howling monkeys in the marajó archipelago, pará, Brazil

I review the distribution of genus Alouattain the Marajó Archipelago based on geographic variation of pelage color patterns. Specific differences exist among the labeled specimens at Museu Paraense Emílio Goeldi, which are confirmed via new field information from wild populations. The animals from Marajó, Caviana, and Mexiana Islands possess color patterns of Alouatta belzebul,while Alouatta seniculusis confirmed as the species that inhabits Gurupá Island. The high variability of pelage coloration of the animals from Marajó, Caviana, and Mexiana Islands includes all phenotypes formerly designated as different subspecies. The howler populations from these three islands belong to the same subspecies as those populations that inhabit the region of Tucurui dam reservoir (area 4). Finally, it seems that, as also suggested by chromosome studies, A. belzebulprobably comprises different epiphenotypes without taxonomic validity.     

Combining phylogeography and landscape genetics of Xenopipo atronitens (Aves: Pipridae), a white sand campina specialist,to understand Pleistocene landscape evolution in Amazonia

Open vegetation (campinas and campinaranas) associated with white sand patches occurs in the form of islands in a forested matrix throughout the Amazon basin. Bird species restricted to these habitats have patchy distributions, although connectivity may have been influenced by past glacial cycles as a result of the substitution of forest by savanna. Because these landscape changes are a matter of debate in the history of Amazonia, we studied the diversification of Xenopipo atronitens, a white sand specialist, aiming to infer the effects of past climate changes. The split of Xenopipo atronitens from its sister species, Xenopipo uniformis, may be related to Tepuis erosion and retreat of escarpments during the Miocene, or to a dispersal event. Compared with birds from terra firme forest, X. atronitens has low genetic structure. Low levels of unidirectional gene flow were found from the Guyana Shield to adjacent areas. Demographic expansion starting approximately 25 kyr BP was detected for some populations and is probably related to the Last Glacial Maximum and subsequent climate improvement. Landscape genetic analyses indicate that the forested (terra firme) matrix acts as a barrier for the dispersal of X. atronitens. The results of the present study indicate that glacial cycles have deeply influenced Amazonian biogeographical history, demonstrating a complex interaction between forest and nonforest habitats during the Pleistocene. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 60–76.     

Changes of the forest-savanna boundary in Brazilian Amazonia during the Holocene revealed by stable isotope ratios of soil organic carbon

The possibility of ecosystem boundary changes in northern Brazilian Amazonia during the Holocene period was investigated using soil organic carbon isotope ratios. Determination of past and present fluctuations of the forest-savanna boundary involved the measurement of natural ¹³C isotope abundance, expressed as ¹³C, in soil organic matter (SOM). SOM ¹³C analyses and radiocarbon dating of charcoal fragments were carried out on samples derived from soil profiles taken along transects perpendicular to the ecotonal boundary. SOM ¹³C values in the upper soil horizons appeared to be in equilibrium with the overlying vegetation types and did not point to a movement of the boundary during the last decades. However, ¹³C values obtained from deeper savanna and forest soil layers indicated that the vegetation type has changed in the past. In current savanna soil profiles, we observed the presence of mid-Holocene charcoals derived from forest species: fire frequency at that time was probably greater, and more extensive savanna may have resulted. Isotope data and the presence of these charcoals thus suggest that the forest-savanna boundary has shifted significantly in the recent Holocene period, forest being more extensive during the early Holocene than today. During the middle Holocene, the forest could have strongly regressed, and fires appeared, with a maximum development of the savanna vegetation. At the beginning of the late Holocene, the forest may have invaded a part of this savanna, and fires occurred again.     

Using Soil 13C to Detect the Historic Presence of Schizachyrium scoparium (Little Bluestem) Grasslands on Martha's Vineyard

Abstract We used differences in soil carbon δ¹³C values between forested sites and grasslands dominated by the C₄ grass Schizachyrium scoparium (little bluestem) to detect the presence of former grasslands in the historical landscape of the coastal sand plain of Martha's Vineyard, Massachusetts, U.S.A. Soil δ¹³C was measured at (1) sites with long‐term forest or grassland vegetation and (2) sites with known histories where forest vegetation invaded grassland and where forest converted to grassland. The δ¹³C of soil under long‐term grassland was –24.1‰ at 0 to 2 cm depth and –23.4‰ at 2 to 10 cm and was enriched by 3.4‰ and 2.8‰ compared with soil under long‐term forest. In forests that invaded grasslands dominated by S. scoparium, soil δ¹³C decreased as C derived from trees replaced C from S. scoparium. This decline occurred faster in surface soils and in the light soil organic matter fraction than in the mineral soil. In forests that converted to grasslands, soil δ¹³C increased and the rate of increase was similar in surface and mineral soil and in the different soil organic matter fractions. Rates of change indicated that soil δ¹³C could be used to detect changes in vegetation involving the presence or absence of S. scoparium during the last 150 years. Application of this model to a potential grassland restoration site on Martha's Vineyard where the landscape history was not known indicated that the site was previously unoccupied by S. scoparium during this time. The δ¹³C of surface mineral soil can be useful for detecting the presence of historic S. scoparium grasslands but only in the period well after European settlement of these coastal sand plain landscapes.     

Secondary forest growth deviation from chronosequence predictions in central Amazonia

Nearly all published rates of secondary forest (SF) regrowth for Amazonia are inferred from chronosequences. We examined SF regrowth on abandoned pastures over a 4‐year period to determine if measured rates of forest recovery differ from chronosequence predictions. We studied the emergence, development and death of over 1300 stems in 10 SFs representing three age classes (<1–5, 6–10 and 11–14 years old). Mean tree biomass accumulation in both the <1–5 and 6–10 years old (4.4 and 5.7 Mg ha⁻¹ yr⁻¹, respectively) abandoned pastures was lower than predicted and deviated significantly (57% and 41%) from rates estimated from the chronosequence. The older SFs, with a mean growth rate of 9.9 Mg ha⁻¹ yr⁻¹ followed the rate predicted by the chronosequence. Understocking was the primary cause of low biomass recovery rates in the youngest forests; although the youngest stands had a diameter at breast height increment three times the oldest stands, the youngest stands lacked sufficient density to cumulatively produce high biomass accumulation rates. Four years of measurement indicated that the youngest stands had developed 59% of the stems measured in the older stands during the same time period. The 6–10‐year‐old stands were rapidly self‐thinning and approached stem density values measured in the same aged stands at the onset of the study. Mortality was high for all stands, with 54% of the original stems remaining after 4 years in intermediate‐aged stands. The forests were dominated by the tree Vismia, which represented 55–66% of the biomass in all stands. The Vismia share of the biomass was decreasing over time, with other genera replacing the pioneer. Our measured rates of regrowth indicate that generalized estimates of forest regrowth through chronosequence studies will overestimate forest regrowth for the youngest forests that were under land use for longer time‐periods before abandonment. Certified Emission Reductions under the Clean Development Mechanism of the Kyoto protocol should consider these results when predicting and compensating for carbon sequestered under natural forest management.     

Low Primate Diversity and Abundance in Northern Amazonia and its Implications for Conservation

Large areas of the Rio Negro basin in Amazonia are covered by continuous tracts of tropical forest, but have few primate species. This is anomalous considering the general relationship between area and number of species. One possibility is that much of the forest is unsuitable habitat for most primates and the area of suitable habitat is much less than the forested area. This has important consequences for the design of reserves and predictions of the consequences of climate change, which tend to be based on broad categories based on satellite images, and not on information of species distributions within those broad categories. The study was conducted through diurnal and nocturnal line‐transect surveys in the Biodiversity Research Program 25‐km² permanent grid in Viruá National Park, which has vegetation associations typical of much of northern Amazonia. The highest primate diversity and abundances occurred in tall terra firme forests (58%), whereas inundated forests and scrublands, which cover 42 percent of the survey grid and 90.8 percent of the Viruá National Park, have virtually no primates. This suggests that parks and reserves in northwestern Amazonia will have to be very large to maintain viable populations of most primates and their ecological interactions, and that very broad habitat categories are not sufficient to make predictions about actual and future suitability of areas for primate conservation.     

Anthropogenic disturbance and the formation of oak savanna in central Kentucky, USA

Aim To deepen understanding of the factors that influenced the formation of oak savanna in central Kentucky, USA. Particular attention was focused on the link between historical disturbance and the formation of savanna ecosystem structure. Location Central Kentucky, USA. Methods We used dendrochronological analysis of tree‐ring samples to understand the historical growth environment of remnant savanna stems. We used release detection and branch‐establishment dates to evaluate changes in tree growth and the establishment of savanna physiognomy. We contrasted our growth chronology with reference chronologies for regional tree growth, climate and human population dynamics. Results Trees growing in Kentucky Inner Bluegrass Region (IBR) savanna remnants exhibited a period of suppression, extending from the establishment date of the tree to release events that occurred c. 1800. This release resulted in a tripling of the annual radial growth rate from levels typical of oaks suppressed under a forest canopy (< 1 mm year^?1) to levels typical of open‐grown stems (3 mm year^?1). The growth releases in savanna trees coincided with low branch establishment. Over the release period, climatic conditions remained relatively constant and growth in regional forest trees was even; however, the growth increase in savanna stems was strongly correlated with a marked increase in Euro‐American population density in the region. Main conclusions Our data suggest that trees growing in savanna remnants originated in the understorey of a closed canopy forest. We hypothesize that Euro‐American land clearing to create pasturelands released these trees from light competition and resulted in the savanna physiognomy that is apparent in remnant stands in the IBR. Although our data suggest that savanna trees originated in a forest understorey, this system structure itself may have been a result of an unprecedented lack of Native American activity in the region due to population loss associated with pandemics brought to North America by Euro‐Americans. We present a hypothetical model that links human population dynamics, land‐use activities and ecosystem structure. Our model focuses on the following three land‐use eras: Native American habitation/utilization; land abandonment; and Euro‐American land clearance. Ecological understanding of historical dynamics in other ecosystems of eastern North America may be enhanced through recognition of these eras.     

Leaf and Soil Nutrients in a Chronosequence of Second‐Growth Forest in Central Amazonia: Implications for Restoration of Abandoned Lands

Subsistence agriculture, cattle ranching, and periodical land abandonment are common land‐use practices in Amazonia. Because changes in land use affect biogeochemical cycles, secondary forests growing after land abandonment develop at varying speeds and spatial patterns, due in part to varying nutrient dynamics. Leaf and soil nutrient concentrations can provide useful information on nutrient cycling processes and strategies of nutrient use by trees that are suitable for introduction to abandoned areas. To understand nutrient dynamics in secondary forests from different regeneration stages, as well as the importance of pioneer species in the regeneration process, we measured the concentration of macronutrients in leaves of three pioneer tree species (Vismia cayennensis, Cecropia sciadophylla, and Bellucia dichotoma) in central Amazon secondary forests. We also measured macronutrients in the topsoil under the trees. We found that type of prior land use, time since abandonment, and number of fire events were significantly correlated with the concentrations of leaf and soil macronutrients, explaining between 10 and 38% of the variation in macronutrient concentrations. The observed patterns suggest that management practices affect the processes involved in N cycling and availability. Of the three pioneer species, C. sciadophylla showed the highest nutrient resorption efficiency and the highest leaf nutritional quality. We suggest that these two features determine a higher potential of C. sciadophylla for natural regeneration and restoration of abandoned lands.