Predation by vertebrates and invertebrates on the seeds of five canopy tree species of an Amazonian forest

We studied the pre-germination loss of seeds to invertebrate and vertebrate seed predators of 5 species of Amazonian trees (Astrocaryum macrocalyx—Palmae; Bertholletia excelsa—Lecithydaceae; Calatola venezuelana—Icacinaceae; Dipteryx micrantha—Leguminosae (Papilionoidae); Hymenaea courbaril-Leguminosae (Caesalpinoidae)). These five species were selected from a large tree flora on several criteria. All possess large (3–10 cm) well-protected seeds that might plausibly be attractive to mammalian seed predators. The reproductive biology of three of the species, or close congeners, had been studied elsewhere in the Neotropics (Astrocaryum, Dipteryx, Hymenaea); one is important to the economy of southeastern Peru (Bertholletia); and one, despite large and apparently edible seeds, appeared to suffer no pre-germination loss to predators (Calatola). We conducted the research in mature forests in the Manu National Park of southeastern Peru where mammal densities are unperturbed by human activities. Densities of adult trees of the five species in our area range from very high (>30 per ha: Astrocaryum) to very low (1 per ha: Hymenaea).Loss of seeds to all causes, and to mammalian seed predators in particular, was determined for seeds placed in 2-square meter mammal exclosures and in open controls located at 10 m (near) and 50 m (far) from a large mature individual of the target species (with minor variations in the design for Astrocaryum and Calatola). The exclosures were of two types: impermeable—designed to exclude all mammals, but not invertebrate seed predators, and semipermeable—designed to admit small (<500 g), but not large mammals. Experimental and control plots were stocked with apparently viable seeds during the dry-wet transition period (October–November) and scored one year later.A significant distance effect (higher predation near vs far from a large conspecific adult) was found in only one of the species (Astrocaryum), the only one to be attacked with high frequency by invertebrate seed predators. The absence of any detectable distance effect attributable to mammals suggests that mammals, over the course of a year, thoroughly search the forest floor for seeds. Invertebrates may thus be responsible for most pre-germination distance (density) effects. With respect to the treatments, we found three qualitatively distinct results: seeds of three species (Astrocaryum, Bertholletia, Dipteryx) were significantly protected by the impermeable, but not semipermeable exclosures, implicating small mammals in seed loss; the seeds of one species (Hymenaea) were significantly protected by exclosures of both types, implicating large mammals; and the seeds of one species (Calatola) exhibited 100% survival, whether or not protected by exclosures.The importance of large mammals as seed predators is generally underestimated in these experiments because semipermeable exclosures may serve as foraging reserves for small mammals. Finally, we noted no relationship between the intensity of mammalian seed predation (as suggested by the survival of unprotected seeds) and the abundance of adults of the five species in the environment. The diversity of results obtained for the five species reveals that large-seeded tropical trees may display a wide range of demographic patterns, and points to the likely importance of post-germination bottlenecks in the population biology of many species, even those that may experience severe pre-germination seed loss.     

Effects of grazing exclusion on species composition in high-altitude grasslands of the Central Alps

Our study examines the effects of grazing exclusion on low-productive subalpine and alpine grasslands of the Central Alps (UNESCO Biosphere Park Gurgler Kamm, Obergurgl, Austria). A long-term exclusion experiment was established in 2000 in the subalpine, the lower, and the upper alpine zone. With exception of the subalpine zone, domestic herbivores have been grazing during the whole growing season. In grazed and exclosure plots species frequencies were recorded for 7 years. We analysed exclosure effects on species number, community composition, life forms, and functional groups.Species richness did not decrease significantly within the exclosures, but changes in species composition occurred in each zone, although some were transitory in nature. The dynamic trends of the plots were significantly explained by the ‘treatment×year’ effect along the whole altitudinal gradient, but the effects decreased considerably with altitude. In the subalpine and upper alpine exclosures, stress-tolerators, species of low or no nutritive value, and mosses showed a decreasing trend, whereas tall grasses (subalpine exclosures), competitors, and species with high or medium nutritive values (lower alpine exclosures) tended to increase.Overall, our 7-year study revealed that several functional groups reacted to grazing, according to our main expectations. We suggest that these effects will intensify in the long term.     

Species composition and diversity of small Afromontane forest fragments in northern Ethiopia

In the highlands of northern Ethiopia, remnants of the original Afromontane forest vegetation are largely restricted to church yards and other sacred groves in a matrix of cropland and semiarid degraded savanna. To assess the potential for natural forest regeneration, species composition and diversity of all forest fragments (10) in a study area of 13,000 ha were analyzed in relation to environmental and soil variables. Using a random design and a density of approximately one plot per 2 ha in all fragments, 31 20×20 m plots were sampled. Indicator species analysis and MRPP tests yielded five communities representing two forest types and one degraded savanna habitat. The forest fragments had a species-poor tree and shrub community in which plots were rather homogeneous and most species abundant. NMDS and analysis of variance indicated that a topographical gradient correlated to soil phosphorus, soil depth, stoniness and the proximity to the river system explained the major differences in species composition and separated moist and dry Afromontane forest communities. The grazing intensity further partitioned the habitats. Present communities and their environmental correlates indicate that the secondary climax forest in the area probably consisted of dry Afromontane forest interlaced by broad strips of moist Afromontane forest along rivers and streams and not a continuous, mono-dominant Juniperus forest as is often presumed. Negative effects of the degraded matrix on forest fragments increased with decreasing patch area and increasing shape irregularity. Nevertheless, all remaining fragments are important for their role in the landscape ecology of the region as refuges and species pools and should be protected and managed accordingly. If seed dispersal from forest fragments into exclosures and subsequent tree recruitment are both successful, the vegetation type most likely to establish is Afromontane savanna woodland, and if managed properly, eventually dry Afromontane forest may arise. Increasing the size of small patches and placing forest plantations and exclosures in the vicinity of small forest fragments is expected to yield the most immediate results. This approach may increase the likelihood of patch colonisation by frugivorous forest birds and thus foster the regeneration of native woody species.     

Land rehabilitation and the conservation of birds in a degraded Afromontane landscape in northern Ethiopia

The few remaining Afromontane forest fragments in northern Ethiopia and the surrounding degraded, semiarid matrix form a habitat mosaic of varying suitability for forest birds. To evaluate the effect of recent land rehabilitation efforts on bird community composition and diversity, we studied bird species distributions in ten small forest fragments (0.40–20.95 ha), five grazing exclosures (10-year-old forest restoration areas without wood extraction and grazing livestock) and three grazed matrix sites during the rainy season (July–October 2004) using 277 one-hour species counts. Based on the distribution pattern of 146 bird species, sites were assigned to one of three bird communities (birds of moist forest, dry forest or degraded savanna), each occupying a well-defined position along an environmental gradient reflecting decreasing vegetation structure and density. All three communities were representative of the avifauna of Afrotropical Highland open forest and woodland with a high proportion of invasive and competitive generalist species (31%). Apart from these, exclosures shared more species with forest fragments (20%) than did the grazed matrix (5%), indicating local ecosystem recovery. By increasing habitat heterogeneity, exclosures have the potential to enhance landscape connectivity for forest birds and are, therefore, an effective instrument for conserving species in a fragmented landscape. However, 52 bird species (36%) occurred exclusively within forest patches and many forest birds that use exclosures are unlikely to maintain viable populations when forest fragments disappear, particularly as forest fragments may be a critical resource during the hot dry season. This highlights the high conservation value of small isolated forest fragments for less tolerant, forest-limited and/or biome-restricted species.     

Time-scale effects in the interaction between a large and a small herbivore

In the short term, grazing will mainly affect plant biomass and forage quality. However, grazing can affect plant species composition by accelerating or retarding succession at longer time-scales. Few studies concerning interactions among herbivores have taken the change in plant species composition into account. In a salt-marsh system, the long-term effects of exclusion of a large herbivore (cattle) on the abundance of a small herbivore (hare) were studied. Excluding cattle grazing for 30 years resulted in large changes in vegetation composition. In general, the cover of tall-growing species increased in the absence of cattle grazing. These long-term changes negatively affected hare grazing intensity. Hares preferentially fed on Festuca rubra and negatively selected tall growing plants, such as Elymus athericus, both in cattle-grazed and long-term ungrazed areas. However, the intensity of hare grazing was not related to the cover of F. rubra. The cover of tall-growing plants (E. athericus, Atriplex prostrata and Juncus maritimus) appeared to be the best predictor and hare grazing intensity decreased sharply with an increase of the cover of tall plants. When cover of tall plants did not increase, hare grazing intensity was not affected. The study shows that the time-scale of the experiment is of prime importance in studying interactions between herbivores. Species that do not seem to influence the abundance of one another or are competing for the same resources on a short time-scale might well be facilitating each other when looking at larger time-scales while taking plant species replacement into account.     

Winners and losers in a long-term study of vegetation change at Moor House NNR: Effects of sheep-grazing and its removal on British upland vegetation

We analysed data collected between 1954 and 2000 from nine long-term experiments designed to assess the effects of sheep-grazing versus no-sheep-grazing at Moor House NNR, an Environmental Change Network site. The experiments were set up between 1954 and 1972 across a range of vegetation types typical of much of upland Britain. Data from this type of experiment are often difficult to analyse and we describe the procedures undertaken to prepare the data for analysis. We fitted the resultant data to the British National Vegetation Classification and used ordination techniques to assess the relative positions of the experiments to each other. Finally, we used Generalized Linear Mixed-effects Modelling within a Bayesian framework to model change of species taxonomic/physiognomic groups through time in both sheep-grazed and ungrazed treatments across all nine experiments; variables included species diversity, Shannon–Weiner index and derived data on occurrence and abundance of species groups based on taxonomy and physiognomy. Hurdle analysis was used to model the species groups; this analysis separated the change through time in both probability of occurrence (binomial distribution) and abundance (Poisson distribution).In the sheep-grazed plots (the “business-as-usual” treatment hence here designated the “control”) there was a reduction in species diversity and a decrease in abundance of vascular plants, grasses, lichens, liverworts and mosses; whereas herbs, sedges and shrubs increased. When probability of occurrence was considered, there was a reduction in number of presences of both lichens and liverworts. Thus, the status quo management of continuous sheep-grazing, even though reduced since 1972, has resulted in a marked change in species composition of these plant communities, with some winners and some losers, but overall they support the concept of biotic homogenization. It is likely that some of these changes were driven by external factors such as elevated atmospheric nutrient deposition. Removal of sheep grazing had some positive benefits; with the herbs, mosses, sedges and shrubs increasing, but faster reductions in grasses and liverworts. Sedges + rushes were stable. It suggested that future monitoring schemes might use either the probability of occurrence of liverworts and lichens, or the abundance of lichens, liverworts, grasses and mosses as sensitive indicators of change in upland Britain.Moreover, during the period that Moor House has been protected as a nature reserve some key plant species groups have declined in spite of reductions in grazing pressure. To reverse this trend requires some form of interventionist management. In order to increase the diversity of vascular plants some form of disturbance will probably be needed, but for bryophytes and lichens this remains a research question. If the results from these small-scale experiments are replicated at the landscape scale a reduction of sheep grazing pressure, for example in rewilding schemes, will have little effect on species composition over a 28–44 year period.     

Vegetation dynamics in a coastal grassland of Hawaii

Vegetation development following goat removal in a tropical grassland with a Mediterranean rainfall seasonality on leeward Hawaii is reported from two permanent monitoring sites. The two sites are equipped with experimental exclosures against feral goats. The sites were monitored annually for a decade (1971–80) for changes in cover by species. The vegetation changes reported relate to both inside- and outside-exclosure developments.Four dynamic categories were recognized among the grassland species following goat removal. These are the decreasers, the increasers and the persistent species. The latter were divided into stable and oscillating persisters. Attention was drawn to the oscillating persisters, as exemplified by the endemic vine Canavalia kauensis, and to the absence of any correlation of its oscillation pattern to the variation in year-to-year rainfall. It was concluded that the oscillation pattern resides in the population it self and can be attributed to its life-cycle phases. It is suggested that synchronized life-cycle dependent death in local populations is an important dynamic phenomenon in certain plant communities. This phenomenon may be considered as separate from phenology and succession, but of similar significance in understanding the processes of vegetation dynamics.Nomenclature of plant names follows St. John (1973).