Deciduous sapling responses to season and large herbivores in a semi‐arid African savanna

Impacts of large herbivores (>5 kg) on woody plants in African savannas are potentially most severe among plants shorter than 1.6 m. It is well established that severe browsing leads to longer shoots, yet prevents saplings from recruiting into adult size‐classes in African savannas. Increased shoot length, indicating faster shoot growth, is often associated with reduced concentrations of tannins and increased nutrient concentrations, suggesting carbon limitation. We hypothesized that, on average, large herbivores suppress stem height or circumference, but increase shoot length. We also hypothesized that if there were concomitant positive effects on nutrients, or negative effects on tannin concentrations, they would be greatest early in the wet season. We sampled saplings of four deciduous woody species (Acacia grandicornuta, Dichrostachys cinerea, Combretum apiculatum and Grewia flavescens) at different stages of the wet season in a large‐scale, long‐term herbivore exclusion experiment in Kruger National Park, South Africa. Plant height, shoot length and stem circumference were generally not adversely affected by large herbivores, suggesting C limitation is rarely present among deciduous saplings in semi‐arid African savannas, allowing them to tolerate browsing. Time since first rainfall emerged as a predominant factor consistently affecting nutrient and tannin concentrations, rather than large herbivores. Nitrogen and phosphorus generally decreased (by 20–50%), while condensed tannin concentration increased (150–350%) during the wet season, except for one species. We postulate that A. grandicornuta is less prone than other species to accumulating tannins during the wet season because of high investment of C in spines. Although nutrient and tannin concentrations were generally not affected by large herbivores, species‐specific responses were evident very early in the wet season, which is when herbivore populations are most likely to be affected by differential forage quality among plants.     

Patch dieback of Colophospermum mopane in a dysfunctional semi‐arid African savanna

Abstract Patch dieback occurred in an almost monospecific Colophospermum mopane (Kirk ex Benth.) Kirk ex J. Léonard woodland in the Northern Province, South Africa, following severe droughts in 1988–1989 and 1991–1992. Discrete patches of dieback and adjacent paired areas of ‘healthy’ vegetation lost an average of 87 and 13% of basal area to mortality, respectively. Whole trees mostly died on ‘dead’ plots, while single‐stem mortality prevailed on ‘live’ plots. Tree mortality decreased with increasing stem number per tree. Patch dieback did not occur on sandy soils. On fine‐textured soils, variation in soil type, topography or slope did not affect dieback. Dieback was influenced by vegetation structure, soil surface condition and soil chemistry. Intense intertree competition, shown by self‐thinning occurring prior to dieback, was a precondition for dieback. Intertree competition had heightened during the 30 years prior to dieback because of an increase in woody cover. Dieback patches had changed from functioning as sinks of water and sediment to sources of these as a result of loss of perennial herbaceous cover, decreased water retention on bared surfaces, and accelerated erosion during 50 years of livestock ranching. Vegetation had thus become increasingly drought‐prone, exacerbated in places by soils with a high sodium concentration. Dieback had occurred because the water requirements of C. mopane could no longer be met during drought years on the dysfunctional patches.     

Plant DNA‐barcode library and community phylogeny for a semi‐arid East African savanna

Applications of DNA barcoding include identifying species, inferring ecological and evolutionary relationships between species, and DNA metabarcoding. These applications require reference libraries that are not yet available for many taxa and geographic regions. We collected, identified, and vouchered plant specimens from Mpala Research Center in Laikipia, Kenya, to develop an extensive DNA‐barcode library for a savanna ecosystem in equatorial East Africa. We amassed up to five DNA barcode markers (rbcL, matK, trnL‐F, trnH–psbA, and ITS) for 1,781 specimens representing up to 460 species (~92% of the known flora), increasing the number of plant DNA barcode records for Africa by ~9%. We evaluated the ability of these markers, singly and in combination, to delimit species by calculating intra‐ and interspecific genetic distances. We further estimated a plant community phylogeny and demonstrated its utility by testing if evolutionary relatedness could predict the tendency of members of the Mpala plant community to have or lack “barcode gaps”, defined as disparities between the maximum intra‐ and minimum interspecific genetic distances. We found barcode gaps for 72%–89% of taxa depending on the marker or markers used. With the exception of the markers rbcL and ITS, we found that evolutionary relatedness was an important predictor of barcode‐gap presence or absence for all of the markers in combination and for matK, trnL‐F, and trnH–psbA individually. This plant DNA barcode library and community phylogeny will be a valuable resource for future investigations.     

Changes in home range size of African lions in relation to pride size and prey biomass in a semi‐arid savanna

Within‐population studies are needed to investigate the extent of, and the factors underlying, intraspecific variation in home range size. We used data from 12 female and 8 male adult lions instrumented with GPS radio‐collars to describe the ranging behaviour of lions in a population from a dystrophic semi‐arid savanna, Hwange National Park, Zimbabwe. We measured prey availability at the home range scale in 2003, 2004, and 2005. For females, home range size increased as pride biomass increased, which is strongly suggestive of expansionism. Once controlled for pride biomass, home range size decreased as prey biomass increased. Pride ranges responded to changes in food abundance on an annual timescale rather than on a seasonal timescale. Female home range size was influenced by the abundance of kudu in the early dry season, whereas it was influenced by buffalo and young elephant abundance in the late dry season. This study shows that female home range size is mainly driven by the size of the pride, but also by prey abundance. Furthermore, female seasonal home range size may be determined, not only by prey abundance, but also by prey dispersion in the landscape. Home range size of males was driven by both prey biomass and the density of female prides.     

Functional differences between summer and winter season rain assessed with MODIS‐derived phenology in a semi‐arid region

Questions: We asked several linked questions about phenology and precipitation relationships at local, landscape, and regional spatial scales within individual seasons, between seasons, and between year temporal scales. (1) How do winter and summer phenological patterns vary in response to total seasonal rainfall? (2) How are phenological rates affected by the previous season rainfall? (3) How does phenological variability differ at landscape and regional spatial scales and at season and inter‐annual temporal scales? Location: Southern Arizona, USA. Methods: We compared satellite‐derived phenological variation between 38 distinct 625‐km² landscapes distributed in the northern Sonoran Desert region from 2000 to 2007. Regression analyses were used to identify relationships between landscape phenology dynamics in response to precipitation variability across multiple spatial and temporal scales. Results: While both summer and winter seasons show increases of peak greenness and peak growth with more precipitation, the timing of peak growth was advanced with more precipitation in winter, while the timing of peak greenness was advanced with more precipitation in summer. Surprisingly, summer maximum growth was negatively affected by winter precipitation. The spatial variations between summer and winter phenology were similar in magnitude and response. Larger‐scale spatial and temporal variation showed strong differences in precipitation patterns; however the magnitudes of phenological spatial variability in these two seasons were similar. Conclusions: Vegetation patterns were clearly coupled to precipitation variability, with distinct responses at alternative spatial and temporal scales. Disaggregating vegetation into phenological variation, spanning value, timing, and integrated components revealed substantial complexity in precipitation‐phenological relationships.     

Diversity and abundance of macro‐invertebrates on abandoned cattle kraals in a semi‐arid savanna

Abandoned cattle (Bos taurus) kraals are sources of habitat heterogeneity in dystrophic semi‐arid African savannas with a strong positive effect on soil nutrients and plant productivity. However, little is known regarding how macro‐invertebrate assemblages vary between abandoned kraals and the surrounding savanna matrix. We tested whether herbaceous biomass and basal and aerial covers and soil nutrients have an effect on aboveground and belowground macro‐invertebrate assemblages. Twelve abandoned kraals were contrasted with their paired control plots for soil characteristics, herbaceous productivity, and macro‐invertebrate assemblages in Save Valley Conservancy, Zimbabwe. Abandoned kraals had significantly higher concentrations of soil nitrogen (N), phosphorus (P), potassium (K), and calcium (Ca) as well as herbaceous biomass and basal and aerial covers than control plots. Both aboveground and belowground macro‐invertebrate species richness were higher on abandoned kraals. However, only belowground macro‐invertebrate diversity (Shannon H′ and Hill number 1) was significantly higher on abandoned kraals. Soil nutrients and herbaceous productivity had positive and significant correlations with the dominant taxa (Coleoptera, Hymenoptera, Hemiptera, Isoptera, and Myriapoda) on abandoned kraals. These results add to the growing body of evidence that abandoned kraals exert significant effects on savanna spatial heterogeneity years later, with implications on ecosystem processes and functioning.     

Leaf gas exchange characteristics and water- and nitrogen-use efficiencies of dominant grass and tree species in a West African savanna

Whereas leaf gas exchange properties are important to assess carbon and water fluxes in ecosystems worldwide, information of this type is scarce for savanna species. In this study, gas exchange characteristics of 2 C₄ grass species (Andropogon canaliculatus and Hyparrhenia diplandra) and 2 C₃ tree species (Crossopteryx febrifuga and Cussonia arborea) from the West-African savanna of Lamto (Ivory Coast) were investigated in the field. Measurements were done in order to provide data to allow the parameterization of biochemically-based models of photosynthesis (for C₄ and C₃ plant metabolic types) and stomatal conductance ; and to compare gas exchange characteristics of coexisting species. No systematic difference was found between grass and tree species for reference stomatal conductance, under standard environmental conditions, or stomatal response to incident light or vapour pressure deficit at leaf surface. Conversely, grass species displayed higher water (1.5-2 fold) and nitrogen (2-5 fold) photosynthetic use efficiencies (WUE and NUE, ratio of net photosynthesis to transpiration and leaf nitrogen, respectively). These contrasts were attributed to the CO₂ concentrating mechanism of C₄ plants. When looking within plant life forms, no important difference was found between grass species. However, significant contrasts were found between tree species, Cussonia showing higher NUE and reference stomatal conductance than Crossopteryx. These results stress the need to account for functional diversity when estimating ecosystem carbon and water fluxes. In particular, our results suggest that the tree/grass ratio, and also the composition of the tree layer, could strongly affect WUE and NUE at the ecosystem scale in West African savannas.     

Variable rainfall has a greater effect than fire on the demography of the dominant tree in a semi‐arid Eucalyptus savanna

Rainfall, fire and competition are emphasized as determinants of the density and basal area of woody vegetation in savanna. The semi‐arid savannas of Australia have substantial multi‐year rainfall deficits and insufficient grass fuel to carry annual fire in contrast to the mesic savannas in more northern regions. This study investigates the influence of rainfall deficit and excess, fire and woody competition on the population dynamics of a dominant tree in a semi‐arid savanna. All individuals of Eucalyptus melanophloia were mapped and monitored in three, 1‐ha plots over an 8.5 year period encompassing wet and dry periods. The plots were unburnt, burnt once and burnt twice. A competition index incorporating the size and distance of neighbours to target individuals was determined. Supplementary studies examined seedling recruitment and the transition of juvenile trees into the sapling layer. Mortality of burnt seedlings was related to lignotuber area but the majority of seedlings are fire resistant within 12 months of germination. Most of the juveniles (≤1 cm dbh) of E. melanophloia either died in the dry period or persisted as juveniles throughout 8.5 years of monitoring. Mortality of juveniles was positively related to woody competition and was higher in the dry period than the wet period. The transition of juveniles to a larger size class occurred at extremely low rates, and a subsidiary study along a clearing boundary suggests release from woody competition allows transition into the sapling layer. From three fires the highest proportion of saplings (1–10 cm dbh) reduced to juveniles was only 5.6% suggesting rates of ‘top‐kill’ of E. melanophloia as a result of fire are relatively low. Girth growth was enhanced in wet years, particularly for larger trees (>10 cm dbh), but all trees regardless of size or woody competition levels are vulnerable to drought‐induced mortality. Overall the results suggest that variations in rainfall, especially drought‐induced mortality, have a much stronger influence on the tree demographics of E. melanophloia in a semi‐arid savanna of north‐eastern Australia than fire.     

Population size structure of trees in a semi‐arid African savanna: Species differ in vulnerability to a changing environment and reintroduction of elephants

The size structure of a plant population provides a snapshot of potential population trend and weak inference on past history. Size structure of 24 tree species was sampled in a savanna reserve in order to assess their potential vulnerability to local extirpation following the reintroduction of elephants. Regression of number per size class based on an expected reverse‐J structure for a healthy population was undertaken in order to classify trend of each species. On account of the weak realism of this expected structure in semi‐arid environments, log‐normal mixture models were also fitted to individual observations in order to identify states, whose distribution, size and shape can indicate potential population persistence. Population structure of 12 species approximated a reverse‐J, indicating potential future growth. Although regressions of seven species were not significant, suggesting vulnerable populations, mixture models of all except one showed multiple states, indicating variable regeneration and recruitment over time. A single state of adult plants indicates that Sclerocarya birrea was vulnerable to local extirpation if adult mortality escalates. Some apparently declining populations displayed multiple states of established individuals suggesting temporally varying regeneration and recruitment, attributed mainly to rainfall variation, but which were judged as likely to persist. Poor recent regeneration of most species was attributed to well below‐average rainfall for the preceding decade. Different savanna tree species apparently ensure persistence based on individualistic responses to population perturbations, but persistence of some following the reintroduction of elephants may be threatened if certain size classes experience escalated mortality. Subsequent monitoring has supported this concern.     

Trophy hunting and perceived risk in closed ecosystems: Flight behaviour of three gregarious African ungulates in a semi‐arid tropical savanna

Although being an important conservation tool in Africa, trophy hunting is known to influence risk perception in wildlife species, thus affecting the behaviour and fitness of most targeted species. We studied the effects of trophy hunting on the flight behaviour of impala (Aepyceros melampus), greater kudu (Tragelaphus strepsiceros) and sable (Hippotragus niger) in two closed ecosystems, Cawston Ranch (hunting area) and Stanley and Livingstone Private Game Reserve (tourist area), western Zimbabwe. Using standardized field procedures, we assessed the flight behavioural responses of the three species in two seasons: non‐hunting (December–March) and hunting (April–November) between March 2013 and November 2014. We tested the effect of habitat, group size, sex, season, start distance and alert distance on flight initiation distance using linear mixed models. Habitat, group size sex and alert distance did not have any effect on flight initiation distance for the three species. The three species were more alert and displayed longer flight initiation distances in the hunting area compared with the tourist area. Flight initiation distances for the three species were higher during the hunting season for the hunting area and low during the non‐hunting season. Flight distances of the three species did not differ between the hunting area and the tourist area. We concluded that trophy hunting increased perceived risk of wild ungulates in closed hunting areas, whereas ungulates in non‐hunting areas are less responsive and somehow habituated to human presence. Management plans should include minimum approach distances by tourists as well as establishing seasonal restrictions on special zones to promote species viability. Research aimed at integrating behavioural responses with physiological aspects of target species should be promoted to ensure that managers are able to deal with the behavioural trade‐offs of trophy hunting at local and regional scale.     

Asymmetry in above‐ and belowground productivity responses to N addition in a semi‐arid temperate steppe

Nitrogen (N) enrichment often increases aboveground net primary productivity (ANPP) of the ecosystem, but it is unclear if belowground net primary productivity (BNPP) track responses of ANPP. Moreover, the frequency of N inputs may affect primary productivity but is rarely studied. To assess the response patterns of above‐ and belowground productivity to rates of N addition under different addition frequencies, we manipulated the rate (0–50 g N m^?2 year^?1) and frequency (twice vs. monthly additions per year) of NH₄NO₃ inputs for six consecutive years in a temperate grassland in northern China and measured ANPP and BNPP from 2012 to 2014. In the low range of N addition rates, BNPP showed the greatest negative response and ANPP showed the greatest positive responses with increases in N addition (<10 g N m^?2 year^?1). As N addition increased beyond 10 g N m^?2 year^?1, increases in ANPP dampened and decreases in BNPP ceased altogether. The response pattern of net primary productivity (combined above‐ and belowground; NPP) corresponded more closely to ANPP than to BNPP. The N effects on BNPP and BNPP/NPP (f_BNPP) were not dependent on N addition frequency in the range of N additions typically associated with N deposition. BNPP was more sensitive to N addition frequency than ANPP, especially at low rates of N addition. Our findings provide new insights into how plants regulate carbon allocation to different organs with increasing N rates and changing addition frequencies. These root response patterns, if incorporated into Earth system models, may improve the predictive power of C dynamics in dryland ecosystems in the face of global atmospheric N deposition.     

Decomposition and nutrient release patterns of mistletoe litters in a semi‐arid savanna,southwest Zimbabwe

Nutrient loss from litter plays an essential role in carbon and nutrient cycling in nutrient‐constrained environments. However, the decomposition and nutrient dynamics of nutrient‐rich mistletoe litter remains unknown in semi‐arid savanna where productivity is nutrient limited. We studied the decomposition and nutrient dynamics (nitrogen: N, phosphorous; P, carbon: C) of litter of three mistletoe species, Erianthemum ngamicum, Plicosepalus kalachariensis, and Viscum verrucosum and N‐fixing Acacia karroo using the litter‐bag method in a semi‐arid savanna, southwest Zimbabwe. The temporal dynamics of the soil moisture content, microbial populations, and termite activity during decomposition were also assessed. Decay rates were slower for A. karroo litter (k = 0.63), but faster for the high quality mistletoe litters (mean k‐value = 0.79), which supports the premise that mistletoes can substantially influence nutrient availability to other plants. Nitrogen loss was between 1.3 and 3 times greater in E. ngamicum litter than in the other species. The litter of the mistletoes also lost C and P faster than A. karroo litter. However, soil moisture content and bacterial and fungal colony numbers changed in an opposite direction to changes in the decomposition rate. Additionally, there was little evidence of termite activity during the decay of all the species litters. This suggests that other factors such as photodegradation could be important in litter decomposition in semi‐arid savanna. In conclusion, the higher rate of decay and nutrient release of mistletoe than A. karroo litter indicate that mistletoes play an important role in carbon and nutrient fluxes in semi‐arid savanna.     

Growth responses of an African savanna tree, <Emphasis Type="Italic">Bauhinia thonningii</Emphasis> Schumacher,to defoliation,fire and climate

This study investigated the growth responses to defoliation, fire-exposure and climate factors of a widespread Africana savanna tree, Bauhinia thonningii Schumacher, at a site in central Zambia. Experimental trees (n = 47) were either exposed to fire (n = 12) in the first half of the dry season or protected from fire (n = 35). Some of the fire-protected trees (n = 12) were subjected to artificial defoliation in two consecutive years. Phenological responses (bud break, leaf flush and leaf production) to fire-exposure and defoliation were monitored on permanently marked sample shoots over a 2-year period. Radial tree growth (diameter at 1.3 m above ground) was measured annually over a 7-year period from 1998 to 2005. During the first two years, defoliation and fire-exposure advanced the onset of bud break and leaf flush but fire-exposed trees produced significantly less leaves than did trees protected from fire, probably because scorching caused more severe shoot die-back than is normal. Leaf production was also significantly affected by experimental treatments and their interaction with year. Although treatments had significant short-term effects on radial growth, previous-year growth significantly influenced current-year growth, thereby confirming the existence of autocorrelation in the time-series growth data of B. thonningii. The interaction between previous-year growth and climate factors explained a significant proportion (25–40%, P < 0.001) of the variance in annual tree growth. Variogram models predicted that a 2-year manual defoliation treatment would shorten the longer-term growth cycle while continuous fire-exposure extended the cycle by one year. The results are useful for the management of savanna trees.     

Within‐plant variation in defences in response to simulated herbivory in a semi‐arid southern African savannah

Within‐plant spatial variation in herbivore pressure can induce localized antiherbivory defence responses. We tested this hypothesis by studying branch‐specific responses of Acacia robusta, Dichrostachys cinerea and Ziziphus mucronata to simulated mammalian herbivory. Herbivory was simulated by clipping the terminal shoots (3 cm from tip) of tree branchlets, allowing them one year of regrowth and then comparing their spine length and density and condensed tannins with those of adjacent unclipped branchlets. Condensed tannins concentrations were higher in clipped branchlets than in unclipped branchlets in all three woody species (P < 0.05). Spine length was higher in clipped branchlets than in unclipped branchlets in A. robusta (P < 0.05) but was similar in both D. cinerea and Z. mucronata (P > 0.05). Spine density was double in clipped branchlets as compared to the unclipped branchlets in Z. mucronata (P < 0.05) but was similar in both A. robusta and D. cinerea (P > 0.05). We found evidence of within‐plant variation in condensed tannins concentration and spine length and density in response to simulated herbivory in the three woody species.     

Interspecific variation in the resprouting responses of Acacia species following simulated herbivory in a semi‐arid southern African savannah

Plants have evolved a diverse suite of tolerance traits against herbivory, including compensatory growth, increased photosynthesis and activation of dormant meristems. We studied the responses of five Acacia species to simulated herbivory in a semi‐arid southern African savannah. We clipped terminal shoots of five juvenile Acacia species (Acacia rehmanniana, A. nilotica, A. karroo, A. arenaria and A. gerarrdii) to simulate herbivory. We then determined biomass change after 5 months and also counted the number of resprouts and measured their length and diameter. All clipped shoots produced resprouts, with all the Acacia species compensating for the lost biomass. We found considerable interspecific variation in the compensation for biomass lost to herbivory in the five Acacia species. Resprouts biomass ranged from two times in A. arenaria to four times that removed in A. karroo. Acacia karroo produced many resprouts, while A. arenaria produced very few resprouts (4 vs 15 resprouts). The relationship between the number of resprouts and their growth also varied among the different Acacia species. We conclude that the response of Acacias to herbivory ranges from prolific resprouters (such as A. karroo) to poor resprouters (e.g. A. arenaria).     

Large‐scale environmental flow results in mixed outcomes with short‐term benefits for a semi‐arid floodplain plant community

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Relationship between group dynamics and spatial distribution of African elephants in a semi‐arid environment

Group dynamics related to distribution of African elephants in the semi‐arid environment of Tsavo East National Park, Kenya was studied between 2007 and 2010. I examined the seasonal distribution of lone bulls, bull groups, family units and mixed groups and group size. Lone bulls were widely distributed in the dry season and localized in the wet season, whereas bull groups were localized in both seasons. On average bull group size did not differ with seasons or areas, but larger groups were observed in preferred areas in the dry season. Family units were widely distributed in the dry season and localized in the wet season, whereas mixed groups were localized in both seasons. Although family units and mixed groups tended to be large in all areas in the wet season, large groups were also observed in the dry season. Large family units were widely distributed, whereas large mixed groups were localized. The relationship between group dynamics and distribution was attributed to the combination of social, ecological and poaching factors. This study provided insight into distribution patterns that can be applied to the security of the Tsavo elephant population.     

Community structure of endemic Mimosa species and environmental heterogeneity in a semi‐arid Mexican valley

Abstract. In this paper we analyse six communities with seven Mimosa species in the Tehuacán‐Cuicatlán valley, Mexico. All species are endemic to Mexico and four are endemic to the valley. Mimosa species are found in (1) the ‘matorral xerófilo’ (arid tropical scrub): Mimosa calcicola, M. lacerata, M. luisana, M. polyantha and M. purpusii, and (2) the ‘selva baja caducifolia’ (tropical deciduous forest): M. adenantheroides, Mimosa texana var. filipes. Most of them occur in similar soil environments, while M. polyantha and M. calcicola establish in particular soil conditions and only M. luisana establishes in two different sites showing a wider range of adaptation to soil characteristics. The communities studied include 24 plant families, 51 genera and ca. 70 species (5% of the total flora estimated in the valley). Heterogeneity was found among the communities. Our results point to the replacement of ‘matorral xerófilo’ and ‘selva baja caducifolia’ by ‘matorral espinoso’ (thorny scrub). Thorny species (e.g. Acacia cochliacantha, Mimosa spp.) are becoming the dominant/codominant elements in the communities. Within the communities, Mimosa species have a significant influence on soil pH, organic matter and electrical conductivity values. These species contribute to ameliorate soil nutrient conditions (OM, N_tot, P, Ca, Mg, Na and K contents) as well as other environmental factors (e.g. temperature, shade) under their canopy in degraded plant communities and may serve as ‘resource islands’.