Patterns of seagrass (Posidonia oceanica) flowering in the Western Mediterranean

The intensive reconstructive sampling (1957–2004, 39 localities), a systematic direct observation (1992–2004, 1 locality) and particular direct observations (66 localities) of Posidonia oceanica meadows were analysed together with temporal series of flowering available in the literature (19 localities). This allowed the examination of temporal and spatial variability in annual flowering prevalence (FP, the fraction of meadows flowering in a given year) and of annual meadow flowering intensity (FI, number of inflorescences per shoot) for the period 1979–2004 across the Western Mediterranean, as well as spatial variability of flowering frequency (FF, the fraction of years that a given meadow has flowered) and shoot flowering probability (Pf, fraction of flowering stalks appeared per annual segment). Each year, on an average 17% of the investigated meadows flowered, ranging from 3 to 86% of meadows among the years. The highest annual FP and FI values were obtained in 2003 (FP=0.86 and mean FI=0.23±0.03 inflorescences shoot⁻¹). A secondary peak of FP and mean FI occurred 9 years earlier, in 1994 (FP=0.44 and mean FI=0.08±0.02). Both peaks of flowering occurred after hot summers. Flowering synchrony in particular years across the Western Mediterranean and clines of increased meadow flowering frequency towards the North and East, suggests the existence of large-scale environmental mechanisms controlling the floral induction. On the other hand, meadow FF and Pf were highly heterogeneous among and within the meadows, indicating that local factors also may play a significant role in flowering induction. When flowering, the Western Mediterranean meadows showed an average 0.11±0.02 inflorescences shoot⁻¹, but FI greatly varied among and along the series (from 0.002 to 0.54 inflorescences shoot⁻¹) and decreased significantly with depth but was independent of meadow shoot density and meadow latitude or longitude. The shoot flowering probability was quite low (0.007±0.002 inflorescences shoot⁻¹ year⁻¹) and exponentially increased with shoot age.     

Detritus-Bacteria-Meiofauna interactions in a seagrass bed (Posidonia oceanica) of the NW Mediterranean

The biochemical composition of the sediment organic matter, and bacterial and meiofaunal dynamics, were monitored over an annual cycle in aPosidonia oceanica bed of the NW Mediterranean to test the response of the meiofauna assemblage to fluctuations in food availability. Primary production cycles of the seagrass and its epiphytes were responsible for relatively high (compared to other Mediterranean systems) standing stocks of organic carbon in sediments (from 1.98 to 6.16 mg Cg^–1 sediment dry weight). The biopolymeric fraction of the organic matter (measured as lipids, carbohydrates, and proteins) accounted for only a small fraction (18%) of the total sedimentary organic carbon. About 25% of the biopolymeric fraction was of microphytobenthic origin. Sedimentary organic carbon was mostly refractory (56 to 84%) and probably largely not utilizable for benthic consumers. The biopolymeric fraction of the organic matter was characterized by high carbohydrate concentrations (from 0.27 to 5.31 mg g^–1 sediment dry weight in the top 2 cm) and a very low protein content (from 0.07 to 0.80 mg g^–1 sediment dry weight), which may be a limiting factor for heterotrophic metabolism in seagrass sediments. RNA and DNA concentrations of the Sediments varied significantly during the year. High RNA and DNA values occurred during the microphytobenthic bloom and in correspondence with peaks of bacterial abundance. Bacteria accounted for a small fraction of the total organic carbon (0.65%) and of the biopolymeric organic carbon (4.64%), whilst microphytobenthos accounted for 3.79% of total organic carbon and for 25.08% of the biopolymeric carbon. Bacterial abundance (from 0.8 to 5.8 × 10⁸ g^–1 sediment dry weight) responded significantly to seasonal changes of organic matter content and composition and was significantly correlated with carbohydrate concentrations. Bacteria might be, in the seagrass system, an important N storage for higher trophic levels as il accounted for 25% of the easily soluble protein. pool and contributed significantly to the total DNA pool (on average 12%). Total meiofaunal density ranged from 236 to 1858 ind. 10 cm^–2 and was significantly related, with a time lag, to changes in bacterial standing stocks indicating that microbes might represent an important resource. Bacterial abundance and biomass were also significantly related to nematode abundance. These results indicate that bacteria may play a key role in the benthic trophic     

Flowering of the seagrass Posidonia oceanica in a north-western Mediterranean coastal area: temporal and spatial variations

Sexual reproduction in Posidonia oceanica (L.) Delile is likely to be highly variable both in space and time. However, there has been little effort to identify at what spatial scale variability exists in the distribution and abundance of reproductive shoots or the magnitude of these variations. Such information is important for elucidating several aspects of the reproductive strategy of this species. Here, by using a reconstructing technique, I estimated the rate at which flowering events occurred over the last decades in three P. oceanica meadows along the coastline of the north-western Mediterranean (Italy). Then, variation in inflorescence frequency (i.e. the percentage of shoots that flowered in a particular year of the total number of shoots present in the sample in that year) was examined on a hierarchy of spatial scales, ranging from a few metres to meadows separated by tens of kilometres over a 3-year period. Separate analyses were also conducted to examine the patterns of spatial variability of several plant characteristics (i.e. shoot age and density of vegetative shoots) that may influence the timing of flowering. In total, ten reconstructed flowering events were recorded during the period 1983–1999. Despite the similar climatic conditions experienced in the three meadows surveyed, no inflorescence scars were found in one meadow. In the other two meadows, the flowering rate fluctuated considerably from year to year (0–26%), and simultaneous flowerings were detected only in 3 years (1993, 1994 and 1995). The pattern of spatial variability in the inflorescence frequency varied among years. An investigation of the flowering events recorded along the north-western Mediterranean coast reveals that in the studied period flowering also occurred across a broad range of meadows. Therefore, flowering could be a far more widespread phenomenon in this sector of the Mediterranean basin than previously thought. Analysis of variance showed significant variation in 1993 at the mean inflorescence frequency on very small scales (metres to tens of metres), whereas variation was not found at any of the scales investigated in 1994 and 1995. Overall, variation in the inflorescence frequency within individual meadows (65–93%) was at least an order of magnitude greater than that among separate meadows (0.5–19%). Patterns of spatial variation in the plant characteristics examined did not reflect those of flowering. It may be hypothesised that internal rhythms of populations, in conjunction with changes in environmental conditions, which are consistent over relatively large spatial (kilometres), are involved in the control of flowering induction. Environmental heterogeneity (in terms of nutrient availability) on a very local scale (metres to tens of metres) and intrinsic factors might affect the distribution of reproductive shoots within P. oceanica meadows.Communicated by R. Cattaneo-Vietti, Genova     

Age and growth of the fan mussel Pinna nobilis from south-east Spanish Mediterranean seagrass (Posidonia oceanica) meadows

The calcitic and aragonitic shell of the fan mussel Pinna nobilis L. contains a record of the environmental changes experienced during its growth. Stable-isotope analyses of oxygen (¹⁸O:¹⁶O) in shell carbonate from the calcitic outer shell-layer have been used to validate the periodicity of clearly defined concentric rings on the aragonitic posterior adductor-muscle scar and to estimate the age and growth of fan mussels growing in Posidonia oceanica (L.) meadows at four locations on the south-east Spanish Mediterranean coast. The stable oxygen-isotope records obtained at intervals along a profile across the shell surface enabled seasonal changes in water temperature to be established, and hence seasonal patterns of shell growth to be inferred. Muscle-scar rings were found to be deposited annually in the shell in the spring and early summer (a period of increasing water temperatures), and represent an interruption in the migration of the posterior adductor muscle along the inner surface of the shell. In small pinnids (<25 cm) accretion of the shell is rapid during the first year, but in the second year it is distinctly slower than at the same time the previous year. This slowing down in growth during the second year coincides with the appearance of the “first” distinct muscle-scar ring, indicating that Pinna nobilis does not form a muscle-scar ring during its first year of shell growth. Maximum growth rates were recorded amongst pinnids from Carboneras, where they achieved a length of 59 cm in 8 yr, whilst those from Aguamarga were estimated to be the oldest (attaining a length of 45 cm in 13 yr). Received: 26 January 1998 / Accepted: 8 October 1998     

Primary production of Posidonia oceanica in the Mediterranean Basin

Primary production of the marine phanerogam Posidonia oceanica (Linnaeus) Delile was measured by lepidochronological analyses at 22 sites in the Mediterranean Sea (Corsica, France, Italy, Sardinia and Turkey), between 1983 and 1992, to determine spatial and temporal variations. Leaf production (blade and sheath) ranged from 310 to 1 540 mg dry wt shoot^–1 yr^–1, depending on site and depth. Rhizome production ranged from 24 to 120 mg dry wt shoot^–1 yr^–1 (6% of average leaf production). At some sites the results obtained by lepidochronological analysis were consistent with earlier results obtained by classic methods (e.g. leaf-marking). While primary production per shoot (mg dry wt shoot^–1 yr^–1) displayed no significant differences between sites, primary production of the P. oceanica meadow (g dry wt m^–2 yr^–1) decreased with increasing depth at all sites studied. This decrease correlated with reduced density of the meadow (number of shoots per m²) with increasing depth. Past primary production was also extrapolated at three sites at the island of Ischia (Italy) for a period of 5 yr in order to determine interannual variations over a period of several years. While major variations were recorded for the surface stations (5 and 10 m depth), production remained stable at the deepest station (20 m depth). Given the large geographical scale of the study (location, depth range), it would appear that while P. oceanica production remains considerable, the values recorded in the literature on the basis of classical analyses (surface stations) represent maxima, and cannot be generalised for meadows as a whole.     

Seasonal changes of benthic bacteria in a seagrass bed (Posidonia oceanica) of the Ligurian Sea in relation to origin,composition and fate of the sediment organic matter

Variations in number and biomass of benthic bacteria were examined in the surface sediments of a Mediterranean seagrass bed [Posidonia oceanica (L.) Delile] in the Gulf of Marconi (northwestern Mediterranean Sea) from 1990 to 1991. The annual dynamics of benthic bacterial density and biomass were compared to changes in elemental (organic C and total N) and biochemical (lipids, proteins, carbohydrates) composition of sediment organic matter, as well as to microphytobenthic biomass, dissolved inorganic nutrients and ATP. Bacterial densities exhibited marked seasonal variations (5.12 to 322.7x10⁸ cells g^-1 sediment dry wt) with highest values in late spring. Bacterial standing stocks (15.8 to 882.33 g C g^-1 of sediment dry wt) were high. Bacterial biomass did not correlate with organic C, total N or to specific biochemical components, but correlated significantly with chlorophyll a, ATP and porewater phosphate concentrations. There is evidence that benthic bacteria were responding to variations of algal biomass. Bacterial biomass accounted, on average, for 30% of total living carbon (calculated on the basis of the ATP concentrations) and 8.4% of total organic carbon.     

Trophic diversity of idoteids (Crustacea, Isopoda) inhabiting the Posidonia oceanica litter

The coexistence of three idoteid species in Posidonia oceanica litter raises the question of trophic diversity and their role in the litter degradation process. Hence, diet composition of Idotea balthica, Idotea hectica and Cleantis prismatica was studied using a combination of gut contents and stable isotopes analysis. Gut content observations indicate that P. oceanica dead leaves are an important part of the ingested food for the three species, although their tissues are constituted of only a small to medium fraction of P. oceanica carbon. Our results also underlined the potential role of these species in the degradation of P. oceanica litter by mechanically fragmenting the litter and by assimilating a small to medium fraction of carbon. Moreover, we showed that there were considerable inter- and intra-specific differences in diet composition. Diet differed between juveniles and adults for I. balthica. Crustaceans are an important food source for adults of I. balthica, while I. hectica indicated a major contribution of algal material. C. prismatica showed an intermediate diet. This trophic diversity is probably one of the factors allowing these species to coexist in the same biotope.     

Influence of hydrodynamic conditions on the production and fate of Posidonia oceanica in a semi-enclosed shallow basin (Stagnone di Marsala, Western Sicily)

An integrated approach using hydrodynamic and transport numerical models, lepidochronology and stable isotope analysis was used to investigate how local hydrodynamic conditions influence the primary production and fate of the seagrass Posidonia oceanica in a Mediterranean semi-enclosed marine system (Stagnone di Marsala). The water mass exchange aptitude of different sectors of the basin was analysed, and data collected were used to select two sectors (colonized by Posidonia oceanica showing the lowest and highest water exchange values) for biological analyses. According to the mean dispersal coefficient differences simulated by the hydrodynamic model, growth rate and primary production of P. oceanica differed between sectors, with average values lower in the central sector where water exchange is lower than in the southern sector. Although P. oceanica coverage and primary production were higher in the southern sector, carbon and nitrogen stable isotope analysis suggests that the transfer of seagrass organic matter to higher trophic levels of the food web was higher in the central sector. The possibility of a link between hydrodynamism, production and fate of organic matter is proposed to explain the observed patterns.     

Influence of hydrodynamic conditions on the production and fate of Posidonia oceanica in a semi-enclosed shallow basin (Stagnone di Marsala,Western Sicily)

An integrated approach using hydrodynamic and transport numerical models, lepidochronology and stable isotope analysis was used to investigate how local hydrodynamic conditions influence the primary production and fate of the seagrass Posidonia oceanica in a Mediterranean semi-enclosed marine system (Stagnone di Marsala). The water mass exchange aptitude of different sectors of the basin was analysed, and data collected were used to select two sectors (colonized by Posidonia oceanica showing the lowest and highest water exchange values) for biological analyses. According to the mean dispersal coefficient differences simulated by the hydrodynamic model, growth rate and primary production of P. oceanica differed between sectors, with average values lower in the central sector where water exchange is lower than in the southern sector. Although P. oceanica coverage and primary production were higher in the southern sector, carbon and nitrogen stable isotope analysis suggests that the transfer of seagrass organic matter to higher trophic levels of the food web was higher in the central sector. The possibility of a link between hydrodynamism, production and fate of organic matter is proposed to explain the observed patterns.     

Stimulation of root formation in Posidonia oceanica cuttings by application of auxins (NAA and IBA)

At the present, the long period (from 3 up to 12 months) to form roots from rhizome cuttings of Posidonia oceanica is the major cause of transplant failure. To promote earlier rooting, the effects of different concentrations (5 and 10 mg/l) of two auxins, namely indole-3-butyric acid (IBA) and α-naphtalen aetic acid (NAA), and the time of collection on rooting of plagiotropic and orthotropic cuttings of P. oceanica were tested. Rooting, survival and growth of cuttings were assessed 1 month after planting in a mesocosm. Results demonstrated that the use of auxin was essential to achieve root initiation within the observation period. Irrespective of plant source and collection time, IBA and NAA (5 mg/l) treatments increased the rooting capacity in cuttings. The highest rooting success (i.e. percentage of the survived cuttings that rooted) obtained was 33%. Results also revealed that the survival rate and percentages of cuttings with leaf growth and emergence of new leaves on their terminal shoot were influenced by the collection time × plant source interaction, but were unaffected by the auxin treatment. Orthotropic cuttings taken in November and February showed the highest survival rate (100%). No differences in survival among months were detected for plagiotropic cuttings. In May and July, more plagiotropic cuttings survived (80±7 and 75±6%) compared to orthotropic cuttings (58±7 and 51±4%). Overall, more plagiotropic cuttings showed leaf elongation compared to orthotropic ones, but the inverse was observed in November. A higher percentage of cuttings with emergence of new leaves was observed in the plagiotropic type as compared to the orthotropic one, and in general February and November were the best months for leaf production (28±4 and 38±7%). Finally, the percentage of cuttings that had changed leaf growth orientation (from orthotropic to plagiotropic) was significantly higher in November (63±13%) compared to the other months and tended to be higher in cuttings treated with the auxins compared to controls. These preliminary findings indicate that exposure to IBA or NAA (5 mg/l) was effective in stimulating rooting in P. oceanica without inhibiting plant growth. Therefore, this simple pre-planting practice should be beneficial in restoration attempts. Based on the trend observed, rhizomes should be prepared preferably in late autumn–winter and then planted into restoration stands to achieve a prompt rooting response and the best survival rate. Clearly, future work is needed to maximize the rooting success of cuttings.     

Some quantitative aspects of seagrass ecology in a coastal lagoon of Mauritius

Seagrass distribution was recorded by snorkel dives on a grid of stations in the waterfront of Club Méditerranée at Mon Choisy-Trou Aux Biches lagoon (NW Mauritius) and subsequently mapped using SURFER 6 computer software. Above-ground (AG) and below-ground (BG) standing biomass in terms of dry weight (DW) and ash-free dry weight (AFDW) as well as shoot density and shoot length were monitored monthly from June1997 to May1998 in a mixed stand of Halodule uninervis and Syringodium isoetifolium (dominant) at a shallow, nearshore station in the lagoon. Measurements of physical and chemical parameters [water temperature, current speed, salinity, pH, dissolved oxygen (DO), nitrate and phosphate concentrations] were made simultaneously, as well as at a reference station (ORE) outside the coral reef. The bottom sediment was analysed for grain size and type composition. Variation patterns were examined and statistical correlations drawn to relate plant performance to the environmental variables measured. The SURFER 6 programme generated a satisfactory contour map of seagrass distribution in the lagoon with a cover range of 0-60%. The densest patches occurred adjacent to the shoreline experiencing weaker water currents (3-13 cm s^-1) rather than near the reef (5-35 cm s^-1), where seagrasses were absent. Sand (0.063-2 mm grain size) constituted 97.2% and 77.6% of the nearshore and near-reef sediment, respectively. The dominant grain types were derived from corals (about 80%) and mollusc shells (about 14%). The recorded range of total standing biomass for H. uninervis was 243.1-468.2 g DW m^-2 (326.9ᇛ.7 g) or 71.7-141.2 g AFDW m^-2 (96.8ᆨ.1 g) and for S. isoetifolium it was 271.7-758 g DW m^-2 (460.4끯.1 g) or 119-220.5 g AFDW m^-2 (155.1ᆮ.5 g), with a maximum biomass increase during September-December in both species. AG:BG biomass ratios were generally <1 and approximated 1 during the warmest months of December-February only. Mean shoot density (1,077-4,364 shoots m^-2 in the overall range of 998-4,428 shoots m^-2) and mean shoot length (10.9-20.8 cm in the overall range of 7-31 cm) in S. isoetifolium were higher than in H. uninervis (1,732-4,137 shoots m^-2 in the overall range of 1,522-4,327 shoots m^-2 and 7.9-13.7 cm in the overall range of 6-20 cm, respectively). Temperature showed strong positive correlations with total AFDW biomass of both species (r=0.755, P<0.01 for H. uninervis; r=0.679, P<0.02 for S. isoetifolium) and with DO (r=0.925, P<0.01). High DO levels (10.7-11.2 mg l^-1) coincided with optimum standing biomass at 27.2°C. Correlations were also strong with shoot density (r=0.881, P<0.01 for H. uninervis; r=0.952, P<0.01 for S. isoetifolium) and shoot length (r=0.752, P<0.01 for H. uninervis; r=0.797, P<0.01 for S. isoetifolium). Under optimal environmental conditions, nutrient inputs from surface run-off or underground freshwater seepage in the lagoon due to heavy rainfall may boost up seagrass biomass, as suggested by positive significant correlations between phosphate levels and AG AFDW biomass (r=0.63, P<0.05 for H. uninervis; r=0.65, P<0.05 for S. isoetifolium) and shoot density (r=0.6, P<0.05 for H. uninervis; r=0.687, P<0.02 for S. isoetifolium). The results generated in this study suggest local seagrass standing biomass is comparable to that reported in monospecific stands from elsewhere. Anthropogenic activities increasingly draw down the resilience of the seagrass beds around Mauritius, and preventative measures are indispensable to achieve coastal ecological stability.     

Temporal variation of metals in the seagrass Posidonia australis and its potential as a sentinel accumulator near a lead smelter

Temporal variation in the concentrations of Cd, Cu, Mn, Ni, Pb and Zn in leaves of the seagrass Posidonia australis was studied at three sites near a lead smelter on the shore of Spencer Gulf, a large hypersaline marine embayment in South Australia, on four occasions from October 1980 to September 1981. Concentrations of Cd, Mn, Pb and Zn of up to 541, 537, 379 and 4241 g g^-1, respectively, were found in leaves collected from the site nearest to the smelter. A substantial temporal variation in the concentrations of these metals in samples from all sites resulted from the combined effect of leaf age and collection strategy. Concentrations of Cd, Cu and Zn in the leaf epibiota were lower than those in the leaves, but the reverse was true for Mn and Ni. The use of seagrass leaves as sentinel accumulators for Cd, Pb and Zn must be based on collections made at the same time of year, or otherwise account for the effect of leaf age on concentrations of the metals in the samples.     

The role of stable isotopes and mercury concentrations to describe seabird foraging ecology in tropical environments

Nitrogen (δ¹⁵N) and carbon (δ¹³C) stable isotopes and contaminants, such as mercury, have been widely used to characterise foraging ecology of temperate and polar seabirds. In this study, for the first time, we used isotopic signatures and mercury levels of feathers and blood of eight tropical seabird species, that forage in a range-gradient between inshore and offshore areas, to describe the foraging habits of a large tropical seabird community (from two neighboring islands of the Seychelles archipelago, western Indian Ocean) during both the breeding and inter-breeding periods. Overall, we found a high overlap in both δ¹⁵N and δ¹³C signatures among species. The high inter-specific overlap in δ¹⁵N values was expected, given the similarities in the diet of the species from this community. However, several unexpected results, such as (1) the consistently higher δ¹⁵N signatures of white terns (Gygis alba), (2) the large variation in inter-specific differences in δ¹⁵N signatures among the sampling groups (season, age, island and tissue) and (3) the consistent low δ¹⁵N values of breeding birds during the northwest monsoon (austral summer), suggest that δ¹⁵N signatures cannot be used as indicators of seabird trophic levels in this community. The high inter-specific overlap in δ¹³C signatures and the absence, during the breeding season, of a δ¹³C gradient that follows the inshore-offshore foraging gradient within the community can be explained by the habitat homogeneity of the Seychelles continental shelf and suggest that birds forage mostly within the limits of this “plateau”. On the other hand, the similarities in δ¹³C values between the breeding and inter-breeding periods in species that are known to show post-breeding dispersal, strongly support the hypothesis of a lack of latitudinal variation in δ¹³C signatures of POM in the central Indian Ocean, and the consequent inaccuracy of δ¹³C values to track seabird movements within this geographic area. Inter-specific differences in mercury levels seem to be related to prey size, while consistent higher mercury concentrations in one of the studied islands suggest different island mercury-backgrounds and possible segregation in foraging areas between the seabirds of the two islands. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Variation in assemblages of fish associated with deep and shallow margins of the seagrass Posidonia australis

Little is known about the fish fauna of deep Posidonia australis seagrass beds in Australia. We investigated associated with the deep and shallow margins of P. australis during two surveys in Jervis Bay, New South Wales. Fish associated with deep and shallow seagrass at each of two areas within a large bed of P. australis were compared over a period of 3 mo (November 1986 to January 1987). This was followed by a larger scale survey in which we compared fish found at the two depths within three large beds of P. australis in Jervis Bay over 2 yr (September 1988 to June 1990). In the smaller scale survey there were marked dissimilarities in relative abhundance and composition of species between samples from deep and shallow P. australis. There were also significant differences in abundance between deep and shallow seagrass on most occasions for the majority of the dominant fish species. As a result of the small-scale survey, we concluded that differences in species composition of fish associated with deep and shallow P. australis may have been due to the presence of species from nearby bare substrate among the sparser leaves of the deep seagrass. The results of the smaller scale survey were not confirmed by the larger survey. The relative abundances of species in samples from deep and shallow P. australis were not dissimilar for all beds. In addition, it was rare for any of the dominant species to have significantly different abundances in deep and shallow seagrass. There was little evidence that the species composition of fish associated with deep P. australis was strongly influenced by the fauna of deep bare substrate. Rather, samples from deep P. australis were far more similar to those from shallow P. australis than to a comparable set of samples collected from deep bare substrata. This study demonstrates how the results and conclusions of small-scale survey work, a feature of many environmental impact asseesments, can be misleading when applied to larger spatial and temporal scales, even within the same bay.     

Central place foraging in wheatears (Oenanthe oenanthe L.): foraging itineraries when feeding nestlings

Summary Parent wheatears (Oenanthe oenanthe L.), foraging to meet their own needs and to provide food for a central place (CP, i.e. the nest), have to make decisions with respect to the configuration of foraging itineraries during round trips in the territory and to the directionality of their movements. These problems were studied in two pairs breeding in an agricultural area in central Swecen. All birds started a round trip by hunting from perches close to the CP and then moving to perches progressively further away in a roughly straight line until the first prey item was loaded. Loaded prey were either delivered singly (single prey loading: SPL) or with other prey (multiple prey loading: MPL). When the bird decided to return with several items to the CP, it abruptly changed the direction of its movements by making a left, right or backward turn and started to visit perches progressively nearer the CP, again following a roughly straight line. The decision to return continue loading was affected by the size of the prey as shown by the fact that prey carried singly was significantly larger than the first item in an MPL for all individuals. The distance to the CP also seemed to affect this decision as prey provisioned singly on average were collected significantly closer to the nest than the first item in an MPL by one pair. Both the size of prey loaded singly and load size of MPL increased with distance from the CP in one pair. The concentrated use of the territory in the other pair made any effect of distance difficult to detect. Great flexibility in foraging decisions was observed in cases when an individual, because of the size of the last prey captured, altered its decision to deliver an MPL and transported an SPL to the nest instead. On the basis of these results we propose a set of rules followed by predators such as wheatears when making decisions about delivering prey to a CP (Fig. 4).     

Comparative feeding ecology of felids in a neotropical rainforest

Summary Diet and habitat use of jaguar, puma, and ocelot, and populations of their mammalian prey, were studied in an undisturbed rainforest in southeastern Peru. Analysis of scats (feces) showed terrestrial mammals to be the chief prey of all three felids, but reptiles and birds were also numerically important in the diets of ocelot and jaguar. Prey diversity is high and the cats evidently take any readily captured vertebrate. For major terrestrial mammal prey of felids, density, biomass, prey/predator ratios, and annual offtake from the study area are estimated. All three cat species seem to hunt by opportunistic encounter of prey. Most mammalian prey species were taken in about the ratios of occurrence, but peccaries were taken by jaguar more often than expected. Most prey of jaguar have a body weight of >1 kg, those of ocelot, 1 kg. Jaguar often used waterside habitats, where they captured caiman and river turtles. Puma did not use these habitats or resources, although the puma prey sample was too small for much inference. The possible effects of felids on study area prey populations are discussed. Large and small cats partition prey at the body weight region where prey switches from low to high reproductive rates.