Research Article
Ground-foraging palm cockatoos (Probosciger aterrimus) in lowland New Guinea: fruit flesh as a directed deterrent to seed predation?
- Christopher E. Filardi, Joshua Tewksbury
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- Published online by Cambridge University Press:
- 27 June 2005, pp. 355-361
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Fruit traits that selectively deter vertebrate seed predators without affecting high-quality seed dispersers are said to exhibit directed deterrence. Directed deterrence has been criticized as being unlikely in natural systems, but has rarely been explicitly tested. We evaluated the potential for directed deterrence to explain the double-layered viscous fruit morphology and fruiting phenology of Terminalia impediens, a common canopy tree endemic to New Guinea. The large fleshy fruits of this tree are consumed and dispersed by cassowaries (Casuarius spp.) and are consumed and killed by palm cockatoos (Probosciger aterrimus). Fruit flesh in this tree species appears to function as a deterrent to seed predation by palm cockatoos; the fruits of T. impediens fall to the ground before ripening, and are avoided by palm cockatoos until after the flesh has rotted off the hard nut. We found that palm cockatoos are able to prey upon seeds from fruits without flesh on the forest floor more efficiently than they can consume seeds from unripe fruit still on the trees. Further, through foraging preference tests, we found that palm cockatoos largely ignore seeds within ripe fruit on the ground, but readily eat the seeds when the fruit flesh has been removed. Cassowaries consume the fruit whole, when ripe, and defecate seeds in piles away from parent trees, where seed predation rates are lower. These results challenge the prevailing view that fleshy fruits evolved in tight synchrony with high-quality seed dispersers and add support to the non-exclusive hypothesis that aspects of fruit fleshiness may also have evolved as a response to seed predation.
Spatial patterns in the understorey shrub genus Psychotria in central Amazonia: effects of distance and topography
- V. F. Kinupp, William E. Magnusson
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- 27 June 2005, pp. 363-374
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We evaluated the effects of topography on the distribution of understorey herbs, shrubs and small trees of the pantropical genus Psychotria (Rubiaceae) in a 10000-ha rain-forest reserve in central Amazonia. As plots were long and thin, and followed altitudinal isoclines, we were able to avoid the trade off between plot size and precision of measurement of topographical variables. The minimum distance between plots (1 km) was sufficient to avoid spatial autocorrelation in topographical variables. However, indices of plot similarity based on species composition were spatially autocorrelated to distances of at least 4 km. Although Multivariate Analysis of Covariance (MANCOVA) indicated significant effects of altitude, slope, and watershed on species composition within plots, topographical variables were generally poor surrogates for species distributions. Differences between eastern and western watersheds within the reserve were not due to distance effects, and most species occurred in both watersheds, indicating that differences in species assemblages between watersheds are determined by ecological factors. Habitat specialization and local density were not clearly associated with rarity. At scales of 1–10 km, both distance and habitat affect the distribution of understorey shrubs of the genus Psychotria, but much of the variation in species abundances remains unexplained.
Do mycorrhizas improve tropical tree seedling performance under water stress and low light conditions? A case study with Dicorynia guianensis (Caesalpiniaceae)
- Moïse Béreau, Damien Bonal, Eliane Louisanna, Jean Garbaye
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- 27 June 2005, pp. 375-381
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We tested the response of seedlings of Dicorynia guianensis, a major timber tree species of French Guiana, to mycorrhizal symbiosis and water limitation in a semi-controlled experiment under natural light conditions. Under well-watered conditions, mycorrhizal colonization resulted in an increase of net photosynthesis, growth and phosphorus uptake. When submitted to water stress, no growth reduction of mycorrhizal seedlings was observed. Mycorrhizal seedlings were more sensitive to drought than non-mycorrhizal ones in terms of carbon assimilation, but not with regard to stomatal closure. In contrast to previous studies on temperate tree seedlings, this result precludes a mycorrhizal effect on the hydraulic properties of this species. Furthermore, our results suggest that below a specific threshold of soil moisture, carbon assimilation of D. guianensis seedlings was decreased by the mycorrhizal symbiosis. This is probably related to the competition between the plant and its host fungus for carbon allocation under low light intensity, even though it did not seem to have a significant effect on mortality in our experiment.
Light-dependent seedling survival and growth of four tree species in Costa Rican second-growth rain forests
- Silvia Iriarte Vivar Balderrama, Robin L. Chazdon
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- 27 June 2005, pp. 383-395
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The dependence of tree seedling survival and growth on light availability was evaluated over 15 mo in three second-growth, wet forest stands (15–18 y old) in north-eastern Costa Rica. Seedlings of four canopy tree species (Dipteryx panamensis, Hyeronima alchorneoides, Virola koschnyi and Vochysia guatemalensis) were planted into four canopy treatments in three replicated stands and in a pasture site, spanning a nearly complete gradient of light availability. Survival and growth of all species increased in response to increasing light availability, but species differed in survival in shaded microsites (6–20% light transmittance) and in growth increments per light increase. Hyeronima showed the highest mortality at low light levels and the highest relative height growth increase per light increase. In contrast, Virola showed high survival at low and moderate light, but showed the lowest leaf area and above-ground biomass per light increase among all species. Dipteryx and Vochysia maintained relatively high rates of survival and growth across the entire light gradient. Hyeronima and Virola showed trade-offs between growth and survival responses to light, unlike Dipteryx and Vochysia. Differences among species in seedling survival at low light may be determined by a variety of physiological and morphological traits that may or may not be mechanistically linked to growth responses.
Responses of seedling transplants to environmental variations in contrasting habitats of Central Amazonia
- Julieta Benítez-Malvido, Miguel Martínez-Ramos, José Luis C. Camargo, Isolde D. K. Ferraz
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- 27 June 2005, pp. 397-406
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In the Central Amazon we investigated whether seedling performance (survival, and relative growth rates in height and leaf numbers) was affected by initial seedling size (height and leaf numbers) in habitats that varied in their degree of human disturbance: cattle pasture, young secondary forest, 1-ha forest fragment and old-growth forest. Additionally, effects of photosynthetically active radiation (PAR), litter standing crop (LSC) and insect herbivory were evaluated 12 mo after transplantation in seedlings from the native canopy trees Chrysophyllum pomiferum, Micropholis venulosa and Pouteria caimito. Seedling performance changed rank across the understorey environment depending on species. Seedlings of Chrysophyllum thrived in all conditions but under high PAR, Micropholis thrived only in intermediate light conditions, whereas Pouteria thrived under high PAR. Effects of initial seedling size, PAR and herbivory after 1 y were specific to species, whereas LSC had no effect on performance. Initially larger seedlings resulted in lower survival for Chrysophyllum and Pouteria. Herbivory affected seedling performance in all species. Negative effects of herbivory were intensified under low PAR. Overall, our results showed that, as seedlings, species of the same family and characteristic of old-growth forests respond differently to the environmental constraints present in contrasting human-disturbed conditions. Larger seedlings may not always present greater tolerance to physical and biotic mortality risks.
Foraging polymorphism in Heliconius charitonia (Lepidoptera: Nymphalidae): morphological constraints and behavioural compensation
- Luis Mendoza-Cuenca, Rogelio Macías-Ordóñez
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- 27 June 2005, pp. 407-415
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Sexes and also within sex phenotypes, frequently differ in morphological traits associated with efficiency and performance in foraging and mating behaviours. In butterflies and other flying animals, phenotypic differences in wing size and traits associated with flight are involved in flight performance and individual fitness, but explorations of links among two or more traits and intrasexual differences are scarce. Foraging patterns were studied in a population of Heliconius charitonia butterflies having three phenotypes (females and two male phenotypes) which differ in their wing morphology and reproductive behaviour. As in previous studies, intersexual differences in foraging patterns were found; more interestingly, intrasexual differences were found between alternative male mating strategies. Using morphological and behavioural data, as well as data from previous flight analyses in Heliconius butterflies, we show that intrasexual differences may be explained by the energetic demands of each phenotype. Energetic expenditure is partially related to phenotypic variability in flight morphology and efficiency, and at least in both male phenotypes, differences may also be related to the energetic demands of alternative mating strategies.
Temporal variation in abundance and diversity of butterflies in Bornean rain forests: opposite impacts of logging recorded in different seasons
- K. C. Hamer, J. K. Hill, N. Mustaffa, S. Benedick, T. N. Sherratt, V. K. Chey, M. Maryati
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- 27 June 2005, pp. 417-425
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We used traps baited with fruit to examine how the temporal variation of butterflies within primary forest in Sabah, Borneo differed between species. In addition, we compared patterns of temporal variation in primary and selectively logged forest, and we tested the hypothesis that selective logging has different recorded impacts on species diversity of adults during the wet monsoon period and the drier remaining half of the year. Species of Satyrinae and Morphinae had significantly less-restricted flight periods than did species of Nymphalinae and Charaxinae, which were sampled mainly during the drier season, especially in primary forest. Species diversity of adults was significantly higher during the drier season in primary forest, but did not differ between seasons in logged forest. As a consequence, logging had opposite recorded impacts on diversity during wetter and drier seasons: primary forest had significantly higher diversity than logged forest during the drier season but significantly lower diversity than logged forest during the wetter monsoon season. The results of this study have important implications for the assessment of biodiversity in tropical rain forests, particularly in relation to habitat disturbance: short-term assessments that do not take account of seasonal variation in abundance are likely to produce misleading results, even in regions where the seasonal variation in rainfall is not that great.
Deep ground fires cause massive above- and below-ground biomass losses in tropical montane cloud forests in Oaxaca, Mexico
- H. Asbjornsen, N. Velázquez-Rosas, R. García-Soriano, C. Gallardo-Hernández
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- 27 June 2005, pp. 427-434
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Although fire is occurring at greater frequencies and spatial scales in the moist tropics, few studies have examined the ecological impacts of fire in tropical montane cloud forest (TMCF). This study, conducted in the Chimalapas region of Oaxaca, Mexico, documents changes in live tree biomass, live fine-root biomass, and fallen and standing dead wood 4 y following deep ground fires occurring in TMCF during the 1997–98 El Niño Southern Oscillation event. Forests growing on two different substrates (metamorphic and sedimentary) and having three different statures (mean canopy heights: 20–30 m, 15–20 m and 4–6 m) were assessed within six paired plots established on adjacent burned and unburned forest sites. Total live tree biomass was 82% and 88% lower for burned TMCF growing on metamorphic and sedimentary substrates, respectively, compared with unburned TMCF. Nearly 100% of the living biomass was killed in elfin TMCF located on exposed sedimentary limestone at the highest elevations. Live fine-root biomass in the upper organic soil horizon of burned TMCF sites was 49% lower on metamorphic substrates and 77% lower on sedimentary substrates compared with unburned sites. The amount of total dead wood was 3- to 14-fold greater in burned forests compared with unburned forests. These results suggest that first-time fires in relatively undisturbed TMCF can cause dramatic changes in live above- and below-ground biomass at levels greatly exceeding values reported for most lowland tropical rain forests. These patterns may be attributed to the slower decomposition rates and thick organic soils typical of TMCF, combined with the relatively fast drainage associated with steep topography and, in some locations, sedimentary limestone-derived substrates.
Fire effects on the spatial patterns of soil resources in a Nicaraguan wet tropical forest
- Brent C. Blair
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- 27 June 2005, pp. 435-444
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Anthropogenic wildfires are becoming increasingly frequent in wet tropical forests. This trend follows that of other anthropogenic disturbances, which are now acute and widespread. Fires pose a potentially serious threat to tropical forests. However, little is known about the impact of unintended forest fires on below-ground resources in these ecosystems. This study investigated the influence of fires on the distribution and variability of soil resources on two sets of 50×50-m burned and unburned plots in a Nicaraguan rain forest. Samples were collected at 5-m intervals throughout each plot as well as subsamples at 50-cm intervals. Geostatistical techniques as well as univariate statistics were used to quantify the spatial autocorrelation and variability of selected nutrients (N, P and K), carbon and standing leaf litter. Most variability in this forest was spatially dependent at a scale of 30 m or less. However the average range of autocorrelations varied greatly between properties and sites. Burning altered soil heterogeneity by decreasing the range over which soil properties were autocorrelated. Overall the average patch size (range) for nitrogen was reduced by 7%, phosphorus by 52%, potassium by 60% and carbon by 43%. While phosphorus and leaf litter increased in the burned plots compared to unburned plots, potassium was not different. Nitrogen and carbon did not display a consistent pattern between burning regimes and this may be explained by variation in fire intensity. Leaf litter measurements did not correlate with measured soil nutrients within plots. Observed changes in the burned forest were likely a result of both the intensity of burning and change in vegetative cover between the time of the fires and soil sampling.
Food selection by a hyperdense population of red howler monkeys (Alouatta seniculus)
- Gabriela Orihuela Lopez, John Terborgh, Natalia Ceballos
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- 27 June 2005, pp. 445-450
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We studied diet choice by a generalist herbivore, the red howler monkey (Alouatta seniculus) under conditions of high and normal population density. Densities equivalent to 800–1000 km−2 (roughly 20–40 times normal) occurred in populations trapped on small, predator-free islands in Lago Guri, Venezuela. For three successive years, we studied one such population, a group of six animals living on a 0.6-ha island and compared its feeding ecology to that of two groups living at normal densities on a 190-ha island. The 0.6-ha island supported a total of 351 trees >10 cm dbh of 46 species, whereas >100 species probably occurred within the 16- and 23-ha home ranges of the two large-island howler troops. Small-island howlers were thus predicted to consume fewer resources, in particular less fruit, and to be less selective in diet choice than large-island howlers. As predicted, small-island howlers consumed fewer resources and obtained a smaller fraction of their intake from rare tree species (those contributing <1% of basal area). Small-island howlers consumed less fruit (2% of feeding time vs. 22%) and more foliage (73% vs. 55%) than the large-island groups. Diet breadth of small-island howlers was markedly less than that of their large-island counterparts. Tree species not present on the small island contributed ≥60% of leaf consumption by large-island howlers. Foliage sources preferred by large-island howlers were different in each of 3 years, whereas foliage of the same species of tree consistently ranked first on the small island. Long-term persistence (17 y) of self-perpetuating howler groups on Lago Guri islets at >20 times normal density strongly suggests that food availability does not limit mainland populations.
Seasonal variation in the leaf gas exchange of tropical forest trees in the rain forest–savanna transition of the southern Amazon Basin
- Eduardo Jacusiel Miranda, George L. Vourlitis, Nicolau Priante Filho, Pedro Correto Priante, José Holanda Campelo, George Sanches Suli, Clóvis Lasta Fritzen, Francisco de Almeida Lobo, Shozo Shiraiwa
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- Published online by Cambridge University Press:
- 27 June 2005, pp. 451-460
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The photosynthetic light response of Amazonian semi-deciduous forest trees of the rain forest–savanna transition near Sinop Mato Grosso, Brazil was measured between July 2000 and September 2003 to test the hypothesis that the photosynthetic capacity of trees acclimated to different growth light environments will decline during the dry season. Maximum photosynthesis (Amax) and stomatal conductance (gmax) were significantly higher during the wet season; however, the physiological response to drought was not a clear function of growth light environment. For some species, such as Psychotria sp. growing in the mid-canopy, internal leaf CO2 concentration (Ci) was >30% lower during the dry season suggesting that declines in Amax were caused in part by stomatal limitations to CO2 diffusion. For other species, such as Brosimum lactescens growing at the top of the canopy, Tovomita schomburgkii growing in the mid-canopy, and Dinizia excelsa growing in the understorey, dry season Ci declined by <20% suggesting that factors independent of CO2 diffusion were more important in limiting Amax. Dry-season declines in gmax appeared to be important for maintaining a more consistent leaf water potential for some species (T. schomburgkii and D. excelsa) but not others (Psychotria sp.). These results indicate that while seasonal drought exerts an important limitation on the physiological capacity of semi-deciduous Amazonian forest trees, the mechanism of this limitation may differ between species.
Nutrient cycling and plant–soil feedbacks along a precipitation gradient in lowland Panama
- Louis S. Santiago, Edward A. G. Schuur, Katia Silvera
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- 27 June 2005, pp. 461-470
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This study addresses patterns of nutrient dynamics on a precipitation gradient (1800–3500 mm y−1) in lowland tropical forest with heterogeneous soil parent material, high plant species diversity and large changes in species composition. Mean foliar concentrations of phosphorus, potassium, calcium and magnesium decreased with increasing precipitation, whereas foliar carbon:nitrogen increased with increasing precipitation. Mean foliar nitrogen:phosphorus varied from 16.4–23.8 suggesting that plant productivity at these sites is limited by phosphorus. Total soil nitrogen increased as a function of foliar litter lignin:nitrogen, whereas net nitrogen mineralization rates decreased with increasing lignin:N indicating that as litter quality decreases, more soil nitrogen is held in soil organic matter and the mineralization of that nitrogen is slower. Extractable phosphorus in soil was negatively correlated with foliar litter lignin:phosphorus, illustrating effects of litter quality on soil phosphorus availability. Overall, the results suggest that variation in plant community composition along this precipitation gradient is tightly coupled with soil nutrient cycling. Much of our understanding of effects of precipitation on nutrient cycling in tropical forest is based on precipitation gradients across montane forest in Hawaii, where species composition and soil parent material are constant. Our results suggest that variation in parent material or species composition may confound predictions developed in model island systems.
Resumen: Este estudio trata sobre los patrones de dinámica de nutrientes en un gradiente de precipitación (1800–3500 mm y−1) en un bosque tropical de tierras bajas con material parental heterogéneo, alta diversidad de especies de plantas, y un gran cambio en la composición de especies. Las concentraciones promedio de fósforo, potasio, calcio y magnesio disminuyen con un aumento en la precipitación, mientras que la proporción carbono:nitrógeno aumenta con un aumento en la precipitación. La proporción promedio de nitrógeno foliar:fósforo varía de 16.4 a 23.8, lo cual sugiere que estos sitios tienen limitaciones de fósforo. El nitrógeno total del suelo aumenta como una función de la proporción de lignina foliar de la hojarasca:nitrógeno, mientras que la tasa de mineralización del nitrógeno neto disminuye con un aumento de la proporción de lignina:nitrógeno, lo cual indica que mientras la calidad de la hojarasca disminuye, más nitrógeno del suelo es retenido en la materia orgánica y la mineralización de ese nitrógeno es más lenta. El fósforo extraíble está correlacionado negativamente con la proporción de lignina foliar de la hojarasca:fósforo, lo cual ilustra el efecto de la calidad de hojarasca en la disponibilidad de fósforo del suelo. En resumen, estos resultados sugieren que la variación en la composición de plantas de una comunidad a lo largo de este gradiente de precipitación tiene un efecto considerable en la retroalimentación de los ciclos nutricionales del suelo. Gran parte de nuestros conocimientos sobre el efecto de los gradientes de lluvia en ciclos nutricionales del suelo han sido basados en estudios de gradientes de precipitación de bosques de montaña en Hawaii, en donde tanto la composición de especies y el material parental del suelo son constantes. Nuestros resultados sugieren que la variación del material parental del suelo y la composición de especies pueden confundir las predicciones desarrolladas en sistemas modelos de islas.
Common allometric equations for estimating the tree weight of mangroves
- Akira Komiyama, Sasitorn Poungparn, Shogo Kato
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- 27 June 2005, pp. 471-477
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Inventory data on tree weights of 104 individual trees representing 10 mangrove species were collected from mangrove forests in South-East Asia to establish common allometric equations for the trunk, leaf, above-ground and root weight. We used the measurable tree dimensions, such as dbh (trunk diameter at breast height), DR0.3 (trunk diameter at 30 cm above the highest prop root of Rhizophora species), DB (trunk diameter at lowest living branch), and H (tree height) for the independent variable of equations. Among the mangrove species studied, the trunk shape was statistically identical regardless of site and species. However, ρ (wood density of tree trunk) differed significantly among the species. A common allometric equation for trunk weight was derived, when dbh2H or DR0.32H was selected as the independent variable and wood density was taken into account. The common allometric equations for the leaf and the above-ground weight were also derived according to Shinozaki's pipe model and its extended theory. The common allometric relationships for these weights were attained with given ρ of each species, when DB2 or dbh2 or DR0.32 was selected as the independent variable. For the root weight, the common equation was derived from the allometric relationship between root weight and above-ground weight, since these two partial weights significantly correlated with each other. Based on these physical and biological parameters, we have proposed four common allometric equations for estimating the mangrove tree weight of trunk, leaf, above-ground part and root.
Brief Report
Sarcocystosis of chital (Axis axis) and dhole (Cuon alpinus): ecology of a mammalian prey–predator–parasite system in Peninsular India
- Maithili M. Jog, Rahul R. Marathe, Shantanu S. Goel, Sachin P. Ranade, Krushnamegh K. Kunte, Milind G. Watve
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- 27 June 2005, pp. 479-482
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The question as to whether predators preferentially kill sick or disabled individuals has been addressed by many ecologists working with different predator–prey systems. Rau & Caron (1979) showed that heavily infected moose were more susceptible to hunting. Kruuk (1972) observed that hyenas appeared to select sick animals in the Serengeti. Vorisek et al. (1998) demonstrated that voles infected with a species of Frenkelia were taken more frequently by buzzards. As a broad generalization, wherever prey capture is difficult and involves large energy expenditure a greater proportion of sick animals seems to be captured (Fitzgibbon & Fanshawe 1989, Holmes & Bethel 1972, Temple 1987). In a host–parasite association where the prey species is an intermediate host and the predator is the definitive host, the capture of the prey is often an essential part of the life cycle. Therefore any mechanism which makes the prey susceptible to predation would enhance the parasite's fitness. In such relationships the susceptibility induced by the parasite can be very specific to the predator host (Levri 1998). Freedman (1990) suggested that a mutualistic association between the predator and parasite might exist. A mutualistic relationship can be said to exist between a predator and a parasite if the cost of harbouring the parasite is less than the benefit of greater success in catching the prey. There is perhaps no demonstrated example of such a mutualism in natural populations since it is difficult to weigh the parasite cost against the predation benefit.