Global climate change is predicted to have large impacts on the phenology and reproduction of alpine plants, which will have important implications for plant demography and community interactions, trophic dynamics, ecosystem energy balance, and human livelihoods. In this article we report results of a 3-year, fully factorial experimental study exploring how warming, snow addition, and their combination affect reproductive phenology, effort, and success of four alpine plant species belonging to three different life forms in a semiarid, alpine meadow ecosystem on the central Tibetan Plateau. Our results indicate that warming and snow addition change reproductive phenology and success, but responses are not uniform across species. Moreover, traits associated with resource acquisition, such as rooting depth and life history (early vs. late flowering), mediate plant phenology, and reproductive responses to changing climatic conditions. Specifically, we found that warming delayed the reproductive phenology and decreased number of inflorescences of Kobresia pygmaea C. B. Clarke, a shallow-rooted, early-flowering plant, which may be mainly constrained by upper-soil moisture availability. Because K. pygmaea is the dominant species in the alpine meadow ecosystem, these results may have important implications for ecosystem dynamics and for pastoralists and wildlife in the region.
With biomass densities comparable to large ungulates and megaherbivores, termites play a key functional role in many tropical savanna ecosystems. This study focuses on vegetated termite mounds (termitaria) constructed by the Termitidae species Macrotermes herus. We studied how resource rich termitaria affect graminoid herbs (Poaceae and Cyperaceae), forbs and woody species composition and diversity. The density of termitaria explained 89% of the variation in dense thickets in the area. Fire tolerant Acacia species dominated the open savanna while fire sensitive species like Grewia spp. and the succulent Euphorbia candelabrum were restricted to termite mounds. Termitaria plots had four times the mean number of woody species and supported three times as many forb species as the adjacent savanna. For woody species, both the Shannon-Wiener index and the Shannon evenness index were higher on temitaria than on the savanna. There were no differences for graminoid herbs, except for the Shannon evenness index which was higher on termitaria. Our results indicate that graminoid herb richness peaks at lower productivity levels than trees and forbs in savanna ecosystems, as also recently found in temperate areas.
Thirteen termite mounds and 13 similar-sized control plots were surveyed in central Zimbabwe in order to study large mammalian browsing and vegetation characteristics. The mounds supported almost twice as many tree species as the control plots and the woody vegetation was denser on mounds compared with the woodland plots. Species of woody plants were recorded along with the percentage of branches browsed (cumulative browsing score) by black rhino, Diceros bicornis, elephant, Loxodonta africana and other browsers combined. In addition we measured how the cumulative browsing score on three woody plant species, Acacia nilotica, Colophospermum mopane and Dichrostachys cinerea, which were common both on and off mounds, was related to the distance from mound centre. Both black rhino and elephant cumulative browsing scores were significantly higher on the mound plants compared with the woodland plots. Cumulative browsing score was negatively related to distance from the mound centre for Dichrostachys cinerea, Colophospermum mopane and Acacia nilotica. We propose that termite mound construction in miombo woodland contributes to sustaining populations of megaherbivores and perhaps some woody species in these areas.
Changes in vegetation cover in northern Chobe National Park (Botswana) were assessed using aerial photographs from1962,1985 and1998, with subsequent ground proofing. In addition, cumulative browsing by elephants and the occurrence of ¢re scars were recorded on random vegetation sites within shrubland (n 20) and mixed woodland (n 20). Coverage of woodland vegetation decreased from 60% to 30% between 1962 and 1998, while shrubland vegetation increased from 5% to 33% during the same period. During the study period, woodland has gradually retreated away from the river front. While riparian forest covered a continuous area along the riverfront in 1962, only fragments were left in 1998. We found a signi¢cant decrease in browse use with increasing distance to the Chobe river for Combretum apiculatum, Combretum elaeagnoides, Combretum mossambicense and other woody plants combined (all P < 0.0001). The occurrence of ¢re (P < 0.0001) and basal area (P < 0.0001) were positively related to distance to the river. Elephant browsing occurred on >70% of available stems within 2 km from the river, while less than 20% of the trees had ¢re scars in the same zone. Beyond 7 km from the river, elephant browsing was reduced to >50% of available stems, while more than 50% of the trees had ¢re scars. The density of any of the shrubs was not related to distance to the river neither within shrubland (all P > 0.05) nor within mixed woodlands (all P > 0.05).La densite¨d'aucunarbuste n'e¨tait lie¨e a© la distance jusqu'a© la rivie© re, ni dans la zone arbustive (ensemble P > 0.05) ni dans la zone arbore¨e me¨lange¨e (ensemble P > 0.05).
Summary 1.Top-down regulation of ecosystems by large herbivores is a topic of active debate between scientists and managers, and a prime example is the interaction between elephants Loxodonta africana and trees in African savannas. A common assumption among wildlife managers is that a local reduction in elephant numbers will ultimately allow woodland to self-restore to a desired former state. Such regeneration is, however, dependent on the survival of seedlings of impacted tree species. We conducted a field experiment to investigate seedling predation in the elephanttransformed Chobe riparian woodland of northern Botswana. 2. We planted seedling gardens in (i) complete exclosures that excluded all herbivores except small rodents and invertebrates, (ii) semi-permeable exclosures that excluded ungulates but included primates, lagomorphs, all rodents, gallinaceous birds, etc, and (iii) completely open plots. Seedlings were of two tree species decreasing in the area ( Faidherbia albida and Garcinia livingstonei ) and two that are increasing ( Combretum mossambicense and Croton megalobotrys ). 3. After 9 months, seedling survival ranged from > 75% for all species in the complete exclosure to < 20% for Faidherbia albida in the open plots. Survival of all seedlings except C. megalobotrys declined precipitously in open plots during the dry season when invertebrates are largely dormant but when impalas Aepyceros melampus (locally abundant ungulates) increase the browse components of their diets. 4. Seedling survival in the open plots was negatively related to local impala density but unrelated to that of any other browser. 5. Synthesis and applications . Our findings relate to the current debate about managing elephants to restore southern African savanna landscapes to desired historical states. Various seedling predators, including the ubiquitous impala Aepyceros melampus , regulate the regeneration of trees from seedlings, and our experiments support the hypothesis that tall closed-canopy woodlands originate during episodic windows of opportunity for seedling survival. To artificially recreate such a window would require the decimation of seedling predators as well as elephants, which is impractical at the landscape scale.
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