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.     

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.     

Forest progression modes in littoral Congo, Central Atlantic Africa

Aim To understand the persistence of a forest–savanna mosaic in places where rainfall data suggest that forest take‐over should take place. To study the various modes of forest encroachment, and the role of human activities to hamper it. Location Data were collected on several forest–savanna ecotones in the coastal region of the Republic of Congo. The sites were chosen to illustrate the differing principal modes of forest expansion, corresponding to different levels of anthropic pressure. Methods The study sites were situated on five transects perpendicular to the ecotone (total sampled area: 1.7 ha) and 10 forest clumps in savanna (with diameters from 3 to 20 m). Along the transects botanical identification, diameter measurement and cartography were performed, while leaf area index was measured at a high resolution (every metre) along two of them. Collected data were analysed using a continuous quantification approach, which is much more useful than classical quadrat analysis. Time calibration of progression rates was performed using a simple model of the growth of the characteristic pioneer species, Aucoumea klaineana. Results The two main different modes are reflected in different successional patterns. The edge diffusion is slow (its rate is evaluated to c. 1 m year^?1) and is characterized by a progressive increase in large‐diameter tree density and shade‐tolerant tree density away from the ecotone. Conversely, savanna to forest phase transition by coalescence of clumps exhibits high tree density remnants distributed in established forest. The composition of these remnants is compatible with that of the forest clumps in savannas. Main conclusions Three functional groups of pioneer trees are distinguished: some occupy the edge (edge pioneer), others establish clumps of forest in savanna (clump pioneers) and the longer‐living A. klaineana ensures the transition to ‘mature’ forest. The two different observed patterns (linear edge progression and clump coalescence) can be understood with the use of a model of forest–savanna dynamics, ‘FORSAT’. The two control parameters are the annual rainfall and the frequency of man‐made fires in each savanna. In particular, an increase in the fire frequency can lead to a shift from the coalescence regime to the edge progression one.     

Past vegetation changes in the Brazilian Pantanal arboreal–grassy savanna ecotone by using carbon isotopes in the soil organic matter

Measurements of the organic carbon inventory, its stable isotopic composition and radiocarbon content were used to deduce vegetation history from two soil profiles in arboreal and grassy savanna ecotones in the Brazilian Pantanal. The Pantanal is a large floodplain area with grass-dominated lowlands subject to seasonal flooding, and arboreal savanna uplands which are only rarely flooded. Organic carbon inventories were lower in the grassy savanna site than in the upland arboreal savanna site, with carbon decreasing exponentially with depth from the surface in both profiles. Changes in ¹³C of soil organic matter (SOM) with depth differed markedly between the two sites. Differences in surface SOM ¹³C values reflect the change from C₃ to C₄ plants between the sites, as confirmed by measurements of ¹³C of vegetation and the soil surface along a transect between the upland closed-canopy forest and lowland grassy savanna. Changes of ¹³C in SOM with depth at both sites are larger than the 3–4 per mil increases expected from fractionation associated with organic matter decomposition. We interpret these as recording past changes in the relative abundance of C₃ and C₄ plants at these sites. Mass balances with ¹⁴C and ¹³C suggest that past vegetational changes from C₃ to C₄ plants in the grassy savanna, and in the deeper part of the arboreal savanna, occurred between 4600 and 11 400 BP, when major climatic changes were also observed in several places of the South American Continent. The change from C₄ to C₃, observed only in the upper part of the arboreal savanna, was much more recent (1400 BP), and was probably caused by a local change in the flooding regime.     

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.     

<Superscript>15</Superscript>N natural abundance of fossil peat reflects the influence of animal-derived nitrogen on vegetation

'¹⁵N signatures of fossil peat were used to interpret past ecosystem processes on tectonically active subantarctic Macquarie Island. By comparing past vegetation reconstructed from the fossil record with present-day vegetation analogues, our evidence strongly suggests that changes in the '¹⁵N signatures of fossil peat at this location reflect mainly past changes in the proportion of plant nitrogen derived from animal sources. Associated with uplift above sea level over the past 8,500 years, fossil records in two peat deposits on the island chronicle a change from coastal vegetation with fur and elephant seal disturbance to the existing inland herbfield. Coupled with this change are synchronous changes in the '¹⁵N signatures of peat layers. At two sites ¹⁵N-enriched peat '¹⁵N signatures of up to +17‰ were associated with a high abundance of pollen of the nitrophile Callitriche antarctica (Callitrichaceae). At one site fossil seal hair was also associated with enriched peat '¹⁵N. Less ¹⁵N enriched '¹⁵N signatures (e.g. -1.9‰ to +3.9‰) were measured in peat layers which lacked animal associated C. antarctica and Acaena spp. Interpretation of a third peat profile indicates continual occupation of a ridge site by burrowing petrels for most of the Holocene. We suggest that ¹⁵N signatures of fossil peat remained relatively stable with time once deposited, providing a significant new tool for interpreting the palaeoecology.     

13C content of ecosystem respiration is linked to precipitation and vapor pressure deficit

Variation in the carbon isotopic composition of ecosystem respiration ('¹³C_R) was studied for 3 years along a precipitation gradient in western Oregon, USA, using the Keeling plot approach. Study sites included six coniferous forests, dominated by Picea sitchensis, Tsuga heterophylla, Pseudotsuga menziesii, Pinus ponderosa, and Juniperus occidentalis, and ranged in location from the Pacific coast to the eastern side of the Cascade Mountains (a 250-km transect). Mean annual precipitation across these sites ranged from 227 to 2,760 mm. Overall '¹³C_R varied from -23.1 to -33.1‰, and within a single forest, it varied in magnitude by 3.5-8.5‰. Mean annual '¹³C_R differed significantly in the forests and was strongly correlated with mean annual precipitation. The carbon isotope ratio of carbon stocks (leaves, fine roots, litter, and soil organic matter) varied similarly with mean precipitation (more positive at the drier sites). There was a strong link between '¹³C_R and the vapor saturation deficit of air (vpd) 5-10 days earlier, both across and within sites. This relationship is consistent with stomatal regulation of gas exchange and associated changes in photosynthetic carbon isotope discrimination. Recent freeze events caused significant deviation from the '¹³C_R versus vpd relationship, resulting in higher than expected '¹³C_R values.     

14C transfer between the spring ephemeral Erythronium americanum and sugar maple saplings via arbuscular mycorrhizal fungi in natural stands

We investigated in the field the carbon (C) transfer between sugar maple (Acer saccharum) saplings and the spring ephemeral Erythronium americanum via the mycelium of arbuscular mycorrhizal (AM) fungi. Sugar maple saplings and E. americanum plants were planted together in pots placed in the ground of a maple forest in 1999. Ectomycorrhizal yellow birches (Betula alleghaniensis) were added as control plants. In spring 2000, during leaf expansion of sugar maple saplings, the leaves of E. americanum were labelled with ¹⁴CO₂. Seven days after labelling, radioactivity was detected in leaves, stem and roots of sugar maples. Specific radioactivity in sugar maples was 13-fold higher than in yellow birches revealing the occurrence of a direct transfer of ¹⁴C between the AM plants. The quantity of ¹⁴C transferred to sugar maple saplings was negatively correlated with the percentage of ¹⁴C allocated to the storage organ of E. americanum. A second labelling was performed in autumn 2000 on sugar maple leaves during annual growth of E. americanum roots. Radioactivity was detected in 7 of 22 E. americanum root systems and absent in yellow birches. These results suggest that AM fungi connecting different understorey species can act as reciprocal C transfer bridges between plant species in relation with the phenology of the plants involved.     

Holocene boundary dynamics of a northern Australian monsoon rainforest patch inferred from isotopic analysis of carbon, (14C and δ13C) and nitrogen (δ15N) in soil organic matter

Abstract Soil organic matter (SOM) was sampled from lateritic soil profiles across an abrupt eucalypt savanna–monsoon rainforest boundary on the north coast of Croker Island, northern Australia. Accelerator mass spectrometry dating revealed that SOM that had accumulated at the base of these 1.5 m profiles had a radiocarbon age of about 5000 years. The mean carbon and nitrogen stable isotope composition of SOM from 10 cm deep layers from the surface, middle and base of three monsoon rainforest soil profiles was significantly different from the means for these layers in three adjacent savanna soil profiles, suggesting the isotopic ‘footprint’ of the vegetation boundary has been stable since the mid Holocene. Although there were no obvious environmental discontinuities associated with the boundary, the monsoon rainforest was found to occur on significantly more clay rich soils than the surrounding savanna. Tiny fragments of monsoon rainforest and abandoned ‘nests’ (large earthen mounds) of the orange‐footed scrubfowl, an obligate monsoon rainforest species, occurred in the savanna, signalling that the rainforest was once more extensive. Despite episodic disturbances, such as tropical storm damage and fires, the stability of the boundary is probably maintained because clay rich soils enable monsoon rainforest tree species to grow rapidly and achieve canopy closure, thereby excluding grass and reducing the risk of fire. Conversely, slower tree growth rates, grass competition and fire on the savanna soils would impede the expansion of the rainforest although high rainfall periods with shorter dry seasons may enable rainforest trees to grow sufficiently quickly to colonize the savanna successfully.     

A classic Caribbean algal ridge, Holandés Cays, Panamá: an algal coated storm deposit

Twenty-seven radiocarbon dates of cores recovered from six drill holes indicate that the relief of the ridge on the seaward edge of the Holandés Cays, San Blas, off the Caribbean coast of Panamá, was formed by storm deposits about 2,000 to 2,800 years ago. Although crustose coralline algae are a dominant component of the surface cover on this outer ridge, they played a minor role in the construction of the framework of this bioherm, which therefore cannot be classified as an algal ridge. The framework of the ridge consists dominantly of Agaricia/Millepora rubble that is extensively lithified by micritic submarine Mg-calcite cement. The present-day surface of this area in the Holandés Cays is primarily one of widespread bioerosion with very little indication of substrate accumulation over the past 2,000 years.     

Evidence of a Holocene and continuing recent expansion of lowland rain forest in humid, tropical North Queensland

Abstract. The identification and radiocarbon dating of charcoal collected under tropical rain forest indicated that sclerophyll forests dominated by Eucalyptus occupied parts of the wet tropical lowlands in the Daintree region of North Queensland at least intermittently from 12,000 yr bp until very recently. The results extend the late Pleistocene expansion of pyrophytic, sclerophyll forests which occurred in the upland rain forests to a humid, megathermic coastal lowlands region. Unlike the early Holocene re-expansion of rain forests which occurred generally on the uplands, the sclerophyll forests in the lowland study area were present until at least 1400 yr bp. Changes in coastal geomorphology and coastline positions during the late Quaternary were examined in the study area by superimposing sea levels derived from published curves on sea-bed contours. The results indicate that a very rapid decrease in the extent of the coastal plain occurred during the late Pleistocene. Between 12,000 and 9000 yr bp, 26 km of the coastal plain was submerged and this would have inevitably resulted in concentrations of Aboriginal populations in the area of the present coastline. It is suggested that burning activities by Aborigines in the coastal lowlands were sufficient to reestablish sclerophyll forests during the latter part of the Holocene from approximately 4000 yr bp following a wanner and wetter period which would have been conducive to rain forest re-expansion. Although the evidence suggests that the most recent rain forest recolonization occurred in the study area more than 1000 years ago, the process is still continuing elsewhere in the wet tropical lowlands in North Queensland. The process of eucalypt forest replacement by rain forest may have accelerated since the arrival of Europeans and the concomitant decrease in Aboriginal management.     

Submillennium-scale migrations of the rainforest-savanna boundary in Colombia: C wiggle-matching and pollen analysis of core Las Margaritas

We present the 11?150-cal-yr-long pollen record Laguna Las Margaritas (3°23′N, 73°26′W; 290 m altitude), located at a site sensitive to climatic change near the transition from the Amazonian rainforest to the savanna of the Llanos Orientales in Colombia. In the 10-m-long core nine AMS ¹⁴C bulk samples show ages from 9760±60 to 854±36 BP and provide initial time control. Thirty-one additional AMS ¹⁴C samples of selected macrofossils provide time control from 6250 to 4050 cal BP; for this interval precise time control was obtained by ¹⁴C wiggle-match dating. From 11?150 to 9100 cal BP, grass savanna dominated the landscape while gallery forest along the drainage system was poorly developed. Water availability was lower than today and the length of the dry season longer. From 9100 to 7330 cal BP gallery forest expanded pointing to wetter conditions. From 7330 to 5430 cal BP savanna was increasingly replaced by forest, but with alternating abundance of both vegetation types. From 5430 to 2500 cal BP forest and wooded savanna dominated the western Llanos Orientales suggesting high precipitation rates. Expansion of Mauritia palm forest at 2500 cal BP reflects increasing water availability and stagnant water environments. Increasing savanna between 2500 and 1000 cal BP may represent a combined natural and anthropogenic signal. Development from open savanna to forest during the Middle Holocene is synchronous with a decreasing caloric seasonality, and a southward migration of the Intertropical Convergence Zone, suggesting that the large-scale climatic and vegetational change in the Colombian savannas is precession-forced. High-frequent migration of the savanna-rainforest boundary started around 6000 cal BP and continued at least to 3000 cal BP. Precipitation regimes in northwest and northeast South America seem opposite: dry conditions in the Colombian savanna area seem to reflect a La Niña setting of the climate system; wet conditions reflect an El Niño setting. Δ¹⁴C fluctuations reflect changes of solar activity and we tested the hypothesis that Δ¹⁴C fluctuations correspond to climatic and vegetational change. For the interval 6250 to 4050 cal BP, we applied ¹⁴C wiggle-match dating (WMD), i.e. matching a series of radiocarbon ages in ¹⁴C BP from the sediment, with the dendrochronology-based calibration curve, and we also wiggle-matched the pollen record vs. the Δ¹⁴C record. In this first attempt in a neotropical ecosystem, we could not find unambiguous support that changes in solar activity did trigger climatic and vegetational change in the savannas of the Llanos Orientales.     

Expansion of gallery forests into central Brazilian savannas

Upland tropical forests have expanded and contracted in response to past climates, but it is not clear whether similar dynamics were exhibited by gallery (riparian) forests within savanna biomes. Because such forests generally have access to ample water, their extent may be buffered against changing climates. We tested the long‐term stability of gallery forest boundaries by characterizing the border between gallery forests and savannas and tracing the presence of gallery forest through isotopic analysis of organic carbon in the soil profile. We measured leaf area index, grass vs. shrub or tree coverage, the organic carbon, phosphorus, nitrogen and calcium concentrations in soils and the carbon isotope ratios of soil organic matter in two transitions spanning gallery forests and savanna in a Cerrado ecosystem. Gallery forests without grasses typically show a greater leaf area index in contrast to savannas, which show dense grass coverage. Soils of gallery forests have significantly greater concentrations of organic carbon, phosphorus, nitrogen and calcium than those of savannas. Soil organic carbon of savannas is significantly more enriched in ¹³C compared with that of gallery forests. This difference in enrichment is in part caused by the presence of C₄ grasses in savanna ecosystem and its absence in gallery forests. Using the ¹³C abundance as a signature for savanna and gallery forest ecosystems in 1 m soil cores, we show that the borders of gallery forests have expanded into the savanna and that this process initiated at least 3000–4000 bp based on ¹⁴C analysis. Gallery forests, however, may be still expanding as we found more recent transitions according to ¹⁴C activity measurements. We discuss the possible mechanisms of gallery forest expansion and the means by which nutrients required for the expansion of gallery forest might accumulate.     

Response of a monsoon forest-savanna boundary to fire protection,Weipa, northern Australia

A numerical floristic analysis of samples across a monsoon forest-savanna boundary, from an area that had been actively protected from fire for 15 years, at Weipa, northern Australia, revealed three communities: (i) a monsoon forest with a low closed canopy composed mainly of tree species with extra-Australian tropical affinities and a sparse ground layer; (ii) an ecotone with a distinct closed microphyll shrub layer beneath the open canopy of savanna trees; and (iii) a savanna dominated by Eucalyptus tetrodonta. The development of the ecotone has occurred since fire protection and is of limited extent within the fire protected block. The monsoon forest occurred on soils with significantly higher concentrations of bauxitic pisoliths than the other two communities. Soils under the monsoon forest had significantly higher concentrations of total K, S, C, N, exchangeable K and Ca, and higher pH and electrical conductivity than for soils of either of the other communities. A positive relationship between woody basal area and concentrations of surface soil total P, N, C, exchangeable Ca, CEC and gravel was detected across a 20 m transect from the ecotone community into the savanna. The invasion of monsoon forest seedlings was greatest in the ecotone, with few occurring in the savanna. It appears that the expansion of the monsoon forest requires the development of a layer of shrubs. The mechanism of this facilitation is unclear, although the possible role of nutrient enrichment by the shrubs requires further investigation.     

Seasonal fluctuations of vagile benthos in the uppermost sublittoral of a maritime Antarctic fjord

Quantitative samples of benthos were taken on a stony beach in the maritime Antarctic (Admiralty Bay, King George Island, South Shetland Islands) during a complete annual cycle. The sampled habitat consisted of cobbles lying on sand and gravel in the fringe between the littoral and sublittoral zones; samples were always taken at low tide just below the water line. In this habitat, abundant macroalgal detritus was observed between stones. This stony beach appeared to be very rich in vagile fauna settled between and under stones. Macrobenthos consisted mainly of amphipods (ca. 85% of total number), gastropods (11%) and nemerteans (3%). The abundance of whole macrofauna ranged to over 50,000 ind. m^-2 and its biomass over 600 g m^-2 (FW). Seven species of Amphipoda and four species of Gastropoda were found. Amphipoda were dominated by Gondogeneia antarctica (over 70% of all amphipods) and Paramoera edouardi (over 20%), whereas among gastropods Laevilitorina antarctica prevailed (over 70%). Unexpectedly high abundance and biomass of Amphipoda were observed in the first half of winter (May/July), surpassing otherwise important summer amphipod abundance. The probable reason for this phenomenon could be high autumn abundance of decaying algae on the beach in the tidal zone, providing detritus that is probably the main food source for Amphipoda.     

Stable isotope enrichment (δ15N and δ13C) in a generalist predator (Pardosa lugubris, Araneae: Lycosidae): effects of prey quality

Analysis of the natural variations in stable isotope ratios in animal tissue may be a powerful tool to reveal the trophic position and feeding preferences of generalist predators such as lycosid spiders. In the present study, changes in ¹⁵N/¹⁴N and ¹³C/¹²C ratios in the lycosid spider Pardosa lugubris (Araneae: Lycosidae), fed with prey of different quality, were investigated. Experimental food chains included three trophic levels: prey media, prey organism and predator. In order to analyse the time course of stable isotope enrichment, different life stages of P. lugubris feeding on Drosophila melanogaster were studied. The ¹⁵N content of hatchlings of P. lugubris was significantly lower than that of their mothers, indicating the existence of nitrogen pools with different ¹⁵N signatures in female P. lugubris. With duration of feeding and progressing development, age and body weight, the ¹⁵N content in spiderlings increased. Starvation resulted in ¹⁵N and ¹³C enrichment in P. lugubris. Being fed with prey of different quality resulted in varying patterns of stable isotope enrichment. ¹⁵N but not ¹³C content consistently increased when fed on high quality prey (Heteromurus nitidus, D. melanogaster and a mixed diet consisting of H. nitidus and D. melanogaster). In contrast, low quality prey (Rhopalosiphum padi, Folsomia candida and a mixed diet consisting of H. nitidus, D. melanogaster and F. candida) resulted in deviations from postulated patterns of stable isotope enrichment with ¹⁵N content being similar to that of starving spiderlings. When fed on high quality prey, ¹⁵N enrichment of P. lugubris was close to ca. 3‰. It is concluded that the analysis of variations in the natural abundance of stable isotopes is particularly helpful in revealing the trophic structure of terrestrial food webs in which polyphagous feeders predominate, as is the case in litter and soil.     

Mesozooplankton community structure and grazing impact in the Polar Frontal Zone of the south Indian Ocean during austral autumn 2002

Mesozooplankton community structure and grazing impact were investigated at 13 stations in the Polar Frontal Zone during the second Marion Offshore Variability Ecosystem Study (MOVES II), conducted during April 2002. Total integrated chl- a biomass ranged between 11.17 and 28.34 mg chl- a m^-2 and was always dominated by nano- and picophytoplankton (<20 µm). Throughout the study, small copepods, mainly Oithona similis and Ctenocalanus vanus, numerically dominated the mesozooplankton community, composing up to 85% (range 30-85%) of the total abundance. Grazing activity of the four most abundant copepods ( O. similis, C. vanus, Calanus simillimus and Clausocalanus spp.), constituting up to 93% of total mesozooplankton abundance, was investigated using the gut fluorescence technique. Results of gut fluorescence analyses indicated that Calanus simillimus and Clausocalanus spp. exhibited diel variability in gut pigments with maximum values recorded at night. In contrast, O. similis and Ctenocalanus vanus did not demonstrate diel variation in gut pigment contents. Ingestion rates of the four copepods ranged from 23.23 to 1462,02 ng(pigm.)ind^-1day^-1. The combined grazing impact of the four copepods ranged between 1 and 36% of the phytoplankton standing stock per day, with the highest daily impact occurring at stations occupied in the vicinity of the Antarctic Polar Front (~35.86% at station 23). Among the copepods, O. similis and Ctenocalanus vanus represented the most important consumers of phytoplankton biomass, collectively responsible for up to 89% (range 15-89%) of the total daily grazing impact. Carbon specific ingestion rates of the copepods varied between 42 and 320% body carbon per day.     

Population Density of Chimpanzees and Gorillas in the Petit Loango Reserve,Gabon: Employing a New Method to Distinguish Between Nests of the Two Species

We studied the vegetation and population density of chimpanzees (Pan troglodytes troglodytes) and gorillas (Gorilla gorilla gorilla) in the Petit Loango Reserve, Gabon. The study site consists mainly of primary and secondary forest, and it also includes patches of savanna, inundated forest, and coastal scrub. The density of herbaceous plants of Marantaceae and Zingiberaceae, which serve as important foods for great apes at other study sites, is extremely low. It was very difficult to distinguish between the nests of chimpanzees and gorillas because most gorilla nests were made in trees. We developed a new method to identify the builder of nests via a combination of field observation of nests and fecal samples, and examination of hair samples under a scanning electron microscope. Nests of chimpanzees were mainly in primary forest, and the estimated density (0.78 individual per km ² ) is higher than the average density in Gabon and comparable to the nest density in the Lopé Reserve, Gabon. Nests of gorillas were mainly in secondary forest, and their estimated density (0.21) is similar to the average next density in Gabon but lower than the gorilla nest density in the Lopé Reserve. The low density of herbaceous foods seemed to influence the density of gorillas but not the density of chimpanzees.     

Plants feed ants: food bodies of myrmecophytic Piper and their significance for the interaction with Pheidole bicornis ants

Several species of Piper (Piperaceae) live in symbiosis with Pheidole bicornis (Formicidae-Myrmicinae) on the southern Pacific slope of Costa Rica. These plants produce small single-celled food bodies (FBs) in leaf domatia, formed by the petiole bases and roofing leaf sheaths. In the present study the dependency of ants on FBs of Piper fimbriulatum as a food source was analysed by comparing the natural abundance of ¹³C and ¹⁵N in ants and FBs. Both '¹³C and '¹⁵N values were very similar between FBs and Pheidole bicornis ants but differed substantially between the plant and other ant species. Therefore we suggest that FBs are a main food source for Pheidole bicornis ants. To strengthen this suggestion, the chemical composition of FBs of four myrmecophytic Piper species was analysed, with special emphasis on the nutritional requirements of inhabiting Pheidole bicornis ants. Standard chemical methods were modified and combined to a novel analysis scheme by which all major FB constituents could be quantified from minute [3-10 mg dry mass (DM)] quantities. Piper FBs mainly consisted of lipids (41-48% of DM) and proteins (17-24% of DM). Soluble carbohydrates and amino acids proved to be quantitatively unimportant. N was predominantly stored as soluble protein and, thus, was easily available to the ants. FBs proved to be a high-energy food source (up to 23 kJ g^-1 DM), with a chemical composition that meets well the nutritional needs of the inhabiting ants.     

Fluctuations of macrobenthic populations: a link between climate-driven river run-off and sole fishery yields in the Gulf of Lions

The Rhone river is the most important input to the Mediterranean Sea, responsible for 50% of the primary productivity of the Gulf of Lions. A highly variable amount of 1-23᎒⁶ t year^-1 of terrestrial material is exported to the sea by the Rhone and stocked on the continental shelf for the most part. Soft-bottom communities off the Rhone delta were dominated by polychaetes both in species richness and abundance, and exhibited strong temporal fluctuations mainly related to flooding events. Floods caused pulses of organic matter followed, with different time lags, by peaks of polychaetes. Opportunistic, short-lived species, such as Mediomastus sp. and Aricidea claudiae, exhibited high short-term peaks in density and biomass a few months after flooding events. Conversely, long-lived species, such as Laonice cirrata and Sternaspis scutata, peaked in density and biomass with a time lag of 1-3 years, and their population increase lasted for a few years. The common sole, Solea solea, is a voracious predator of polychaetes which represent >80% of its prey. A positive correlation was found between the mean annual discharge of the Rhone river and the annual commercial landings of S. solea with a time lag of 5 years in the two fishing harbours (Sete and Martigues) located close to the Rhone delta. The long-term increase in food (i.e. polychaete density and biomass) after flooding events might favour the different stages of the sole life cycle, enhancing its population size for several years. Fluctuations of sole fishery yields in the Gulf of Lions could be influenced by climate, as the Rhone river flow is related to the North Atlantic Oscillation that drives precipitation over Western Europe.