Recent papyrus swamp habitat loss and conservation implications in western Kenya

Papyrus Cyperus papyrus swamps are characteristic of many wetlands of tropical Africa. Like most wetland habitats worldwide, they are under human pressure due to harvesting and reclamation for agriculture. Changes in papyrus cover were assessed using aerial photographs at three Important Bird Areas (IBAs) in the Kenyan sector of Lake Victoria. Papyrus area losses of 50% (Dunga), 47% (Koguta) and 34% (Kusa) occurred between 1969 and 2000. The habitat loss and degradation at the sites appeared purposeful, driven by demand for agricultural land and other papyrus products. Cleared papyrus and cultivated areas around all sites increased over the same period. If papyrus habitat loss continues at current rates, papyrus swamps at Dunga and Koguta will disappear by 2020, and Kusa will be reduced to only 19% of its 1969 area. Human population growth around the sites, with concomitant increase in land use activities and papyrus harvesting are the major factors that account for papyrus area reductions. Loss of total habitat is accompanied by deterioration in habitat quality within remaining areas. Papyrus physical structure (height and density) inversely correlate to human disturbances that include footpaths, cutting, burning, grazing and farming. The continued papyrus habitat loss and degradation represents a significant threat to biodiversity conservation particularly for papyrus-specialist birds and other papyrus-reliant species in western Kenya. The observed pattern of papyrus extents and land use changes at all sites provide the site-scale information necessary for papyrus conservation planning. In particular, conservation action is needed most urgently at Dunga and Koguta as they face severe land use pressures.     

Managing the natural capital of papyrus within riparian zones of Lake Victoria,Kenya

The harvesting of natural products such as papyrus (Cyperus papyrus L.), whether for subsistence value or for the production of commodities intended for sale at local markets, contributes to the well-being of riparian peoples around Lake Victoria, Kenya. Serious losses of papyrus wetlands across East Africa have been reported, most of which are attributed to increasing anthropogenic stressors. Recent studies have called for restoration of these wetlands, emphasizing the need for sustainable harvesting strategies to be put in place, although few have provided suggestions as to how this might happen in practical terms and, crucially, with the consent and active participation of local communities as key stakeholders. Here we explore the socioeconomic characteristics of livelihoods based on papyrus, presenting data generated from surveys, interviews and group discussions collected at multiple sites within the Nyando river basin, Kenya. Conceptualizing papyrus stands as living stocks of natural capital, we then outline our proposal for maintaining the provisioning services of this species, without compromising the critical ecohydrological functions of these swamps as land–water buffer zones. Finally we suggest how this approach might be adapted for wider dissemination around Lake Victoria and beyond, motivated by what we believe to be the first reported case of successful papyrus restoration by a local community.     

Primary productivity of papyrus (Cyperus papyrus) in a tropical swamp; Lake Naivasha,Kenya

Papyrus (Cyperus papyrus) standing biomass and the primary productivity of undisturbed and previously harvested areas of papyrus was measured in Lake Naivasha swamp, Kenya. Papyrus culm density in undisturbed swamp was estimated to be 13·1±1·9 culms m⁻² and aerial biomass was 3602 g m⁻². In undisturbed swamp the aerial productivity was 14·1 g m⁻² day⁻¹ while the previously harvested swamp reached a peak of 21·0 g m⁻² after 6 months. The annual aerial production rate of papyrus in Lake Naivasha was estimated to be 5150 g m⁻² year⁻¹. To sustain yields of regularly harvested papyrus swamps, the harvest intervals should exceed 1 year.     

Papyrus swamp development in the Upemba Basin, Zaïre: studies of population structure in Cyperus papyrus stands

The ecology and distribution of papyrus swamps are briefly reviewed together with the ecology, distribution structure and growth characteristics of Cyperus papyrus.
Three age-classes of culm are recognized and girth and density of each are recorded along transects through mree swamps in the Upemba Basin of Zaïre. A regression of culm-unit biomass on culm girth, developed for Ugandan swamps, is used to convert Upemba measurements into biomasses. It is shown that simple measurements, taken at only a single point in time, yield much information on both the structure and the dynamics of papyrus populations.
It is suggested that the standing-crops of such populations are determined by available light, and by local temperature regimes, and are therefore constant, irrespective of site quality (nutrient status), for any particular climatic zone. Culm-duration (life-time) is not significantly influenced by climate and the ratio culms/rhizomes provides a useful indicator of comparative productivity. Unlike community biomass, productivity can be limited by site quality.     

Nitrogenase activity in the papyrus swamps of Lake Naivasha,Kenya

Nitrogenase activity (acetylene reduction activity) was found to occur universally in the Cyperus papyrus swamp in Lake Naivasha. Low rates of acetylene reduction activity (0.9–104.9 nmol C₂H₄ g d.wt. roots^-1 h^-1) were associated with excised roots of C. papyrus but higher rates of activity (89.0–280.4 nmol C₂H₄ g d.wt. roots^-1 h^-1) were associated with intact root systems of the plant. It was estimated that nitrogen fixation associated with young roots alone could supply about 26% of the nitrogen requirements of growing papyrus plants. Acetylene reduction activity in the lake bottom sediments was generally low and associated with adjacent papyrus stands. Plate counts of putative aerobic and facultatively anaerobic N₂-fixing bacteria associated with papyrus roots showed the presence of high numbers of diazotrophs (5.4 × 10⁶ CFU g d.wt. roots^-1). Fewer numbers of N₂-fixing bacteria were detected in the sediments (1.9 × 10³-3.2 × 10⁴ CFU g d.wt. sediment^-1).     

Effect of harvesting on temporal papyrus (<Emphasis Type="Italic">Cyperus papyrus</Emphasis>) biomass regeneration potential among swamps in Winam Gulf wetlands of Lake Victoria Basin,Kenya

This study established for the first time the impact of harvesting on post-harvest papyrus (Cyperus papyrus L.) biomass regeneration potential, with two harvesting regimes compared. Above-ground papyrus biomass was determined. Biomass varied with site. Site had no effect on regeneration potential, but monthly harvesting reduced papyrus biomass regeneration potential among sites. However, seasonal (6-monthly) harvesting did not appear to affect papyrus biomass regeneration potential. Exponential and polynomial trend analyses revealed a consistent downward trend for monthly harvest biomass, and the polynomial trend was more linear (F = 97.913; P < 0.001) than periodic (F = 9.617; P < 0.05). The polynomial trend scenario indicated how papyrus biological dynamics are likely to behave as monthly harvests are repeated. This suggests that regeneration potential is significantly reduced with successive monthly harvest, leading to weak spatial connectivity, papyrus stand fragmentation, and increased landscape patchiness. A 6-month harvest regime can be established to regenerate more biomass between harvests than is currently the case, with positive implications for wetland conservation and carbon sequestration. Papyrus harvesters can be kept off the swamps by establishing a riparian buffer zone of agro forestry trees and shrubs which can substitute for the papyrus as it is left to mature. However, while the information presented is useful for papyrus wetland management strategies, it is recognized that the study period was too short to permit a generalized recommendation.     

Impacts of the C4 sedge Cyperus papyrus L. on carbon and water fluxes in an African wetland

Fluxes of CO₂ and H₂O vapour were measured by eddy covariance from a stand of the C₄ emergent sedge Cyperus papyrus (papyrus), which formed a fringing swamp on the north-west shore of Lake Naivasha, Kenya. The fluxes of CO₂ and H₂O vapour between the papyrus swamp and the atmosphere were large but variable, depending on the hydrology of the wetland system and the condition of the vegetation. These measurements, combined with simulation modelling of annual fluxes of CO₂, show that papyrus swamps have the potential to sequester large amounts of the carbon (1.6 kg C m^–2 y^–1) when detritus accumulates under water in anaerobic conditions, but they are a net source of carbon release to the atmosphere (1.0 kg C m^–2 y^–1) when water levels fall to expose detritus and rhizomes to aerobic conditions. Evapotranspiration from papyrus swamps (E) was frequently lower than evaporation from open water surfaces (E _o) and plant factors have a strong influence on the flux of water to the atmosphere. For the period of measurement E/E_o was 0.36.     

Biomass Allocation in Cyperus papyrus in a Tropical Wetland,Lake Naivasha,Kenya1

The nutrient status of Lake Naivasha, a freshwater lake in southeastern Kenya, has been rising since at least 1982. A potential effect of increases in nutrient supply to the lake's floating papyrus is increasing of the plants’ investment in above–water material and reduction of the amount of energy invested in uptake and storage. Biomass and its allocation between culms, panicles, roots, and rhizomes was measured in 17 sites around the 150–km² lake. Although above–water biomass was greatest in sites closest to the lake's major nutrient inflow, the River Malewa, there was little evidence of corresponding decreases in the biomass of uptake and storage tissues. In August 1995, the average ± SD biomass of papyrus in the lake was 11,540 ± 3020 g/m², with the papyrus containing about 4500 ± 1900 g total carbon/m² and about 100 ± 70 g total nitrogen/m². Plant nitrogen contents did not vary with distance from the main external nutrient supply. Together with low nitrogen concentrations in the plants (0.60 ± 0.26 in culms and 0.99 ± 0.50% in rhizomes), very high carbon to nitrogen ratios (49 ± 20:1) and nitrogen fixation in the rhizosphere explaining only about half of the plants’ nitrogen, papyrus is a likely net sink for nitrogen supplied from the lake's increasingly cultivated watershed. Despite this role, clearance of papyrus in favor of agriculture partly explains the reductions in the area of papyrus within the lake basin from 48 km² in the late 1960s to 14 km² in 1995.     

The potential use of papyrus (Cyperus papyrus L.) wetlands as a source of biomass energy for sub‐Saharan Africa

Four of five people in sub‐Saharan Africa rely on the traditional use of solid biomass, mainly fuelwood, for cooking. In some areas, the current rate of fuelwood consumption will exhaust biomass reserves within the next decade or two. A largely unrecognized source of biomass are tropical wetland ecosystems which have been shown to be some of the most productive ecosystems globally, exhibiting rates of net primary productivity comparable with high‐input, intensively managed agricultural systems. Papyrus (Cyperus papyrus L.) is an emergent sedge with C₄ photosynthesis which is native to the wetlands, river valleys and lakes of central, eastern and southern Africa. The mean standing dry matter of culms and umbels measured at a number of locations throughout East Africa is 38.3 ± 21.6 tDM ha^?1, and the aerial net primary productivity ranges between 25.9 and 136.4 tDM ha^?1 yr^?1. Papyrus vegetation can be harvested by hand and stacked on the rhizome mat for partial air‐drying, and it has been demonstrated that an annual harvesting regime has no negative impacts on long‐term productivity. The use of papyrus as a biofuel for cooking and heating depends on converting it to a suitably combustible form, such as compressed or carbonized briquettes with a calorific value approximately one‐third less than wood charcoal. While papyrus has significant potential as a biofuel, we argue that an integrated management and decision‐making framework for the sustainable utilization of papyrus wetlands is required, in which all ecosystem services including the provision of biomass energy need to be assessed. Sustainability of papyrus wetlands requires management which combines the strength of traditional communal governance and modern legislation to promote its utilization. In this way, local communities can benefit from the inherent advantages of tropical wetlands as very productive ecosystems.     

Offshore refuges support higher densities and show slower population declines of wintering Ruddy Turnstones Arenaria interpres

ABSTRACT

Capsule: Wintering Ruddy Turnstones Arenaria interpres occur in higher densities and their populations decline less on, or close to, offshore refuges than on mainland sites subject to greater levels of human disturbance.

Aim: To compare wintering densities of Ruddy Turnstones and changes in counts across time from sites with differing levels of human disturbance.

Methods: Long-term counts of Ruddy Turnstones (1998/1999 to 2015/2016) were used from 19 sites (two offshore refuges and 17 mainland sites subject to higher levels of human disturbance) in northeast England. No direct measure of human disturbance was available for our mainland sites; instead we used questionnaires (n?=?690) to understand how far people travelled to visit the coast and then used this distance with human population densities in a buffer around each site as a proxy for human disturbance levels.

Results: After controlling for the extent of their preferred habitat at each site (rocky shore) we found: (i) the closer each of the 19 sites was to the nearest offshore refuge the higher the density of Ruddy Turnstones and (ii) bird counts were stable at the two refuge sites, whereas, on average, counts declined at the 17 mainland sites. However, no relationship was found between Ruddy Turnstone counts from 17 mainland sites and human population densities within differing distances from each site (up to 10?km).

Conclusions: Our work suggests that Ruddy Turnstones made greater use of relatively undisturbed areas (offshore refuges) than those subject to greater disturbance by humans (mainland sites). Although the use of refuges and mainland in our study area was not well known, observations from 11 radio-tagged Ruddy Turnstones suggest that individual birds did use both locations. In a broader context, our work concurs with other studies that highlight the need for refuges with limited or no human access.     

Effects of fire on bird diversity and abundance in an East African savanna

Fire is an important determinant of many aspects of savanna ecosystem structure and function. However, relatively little is known about the effects of fire on faunal biodiversity in savannas. We conducted a short‐term study to examine the effects of a replicated experimental burn on bird diversity and abundance in savanna habitat of central Kenya. Twenty‐two months after the burn, Shannon diversity of birds was 32% higher on plots that had been burned compared with paired control plots. We observed no significant effects of burning on total bird abundance or species richness. Several families of birds were found only on plots that had been burned; one species, the rattling cisticola (Cisticola chiniana), was found only on unburned plots. Shrub canopy area was negatively correlated with bird diversity on each plot, and highly correlated with grass height and the abundance of orthopterans. Our results suggest that the highest landscape‐level bird diversity might be obtained through a mosaic of burned and unburned patches. This is also most likely to approximate the historical state of bird diversity in this habitat, because patchy fires have been an important natural disturbance in tropical ecosystems for millennia.     

Papyrus swamps and the respiratory ecology of Barbus neumayeri

Synopsis In seasonally flooding fresh waters, dissolved oxygen and fluctating water levels combine to create strong seasonal changes in habitat availability and dispersal pathways for fishes. In this study we demonstrate how respiratory mode can affect the use of and dispersal through hypoxic papyrus swamps for a small cyprinid fish, Barbus neumayeri, a species which uses aquatic surface respiration (ASR) in response to severe hypoxia. Monthly records of the distribution and relative abundance of B. neumayeri across 28 stations in a papyrus swamp in Uganda were used to examine seasonal patterns of habitat use and movement. The distribution of B. neumayeri was very restricted during the drier months. Most fish were captured in open pools and channels with large areas of open water surface and higher oxygen. Movement of B. neumayeri was limited to short periods during the wet season when peak water levels produced high levels of oxygen and pathways for movement. Three lines of evidence suggest that these patterns relate to its respiratory mode. These include: the positive :relationship between dissolved oxygen and the number of stations used per month, the positive relationship between fish density at dry season stations and dissolved oxygen, and the absence of fish from stations with. little open water surface area during the dry season. Significant differences in the gill morphology between B. neumayeri from the papyrus swamp and those from the main river suggest that papyrus swamps may contribute to the diversification of B. neumayeri by limiting movement and demanding specialization for extremely hypoxic waters.     

Fish communities in the eastern,seasonal-floodplain of the Sudd,Southern Sudan

Annual flooding of the permanent Sudd swamps innundates large areas of Oryza longistaminata grasslands. Fish migration into a southeast section of the seasonal floodplain was led by Clarias gariepinus, Polypterus senegalus and several small species which were intent on spawning. Channa obscura and juveniles of Oreochromis niloticus were prominent among later entrants. From a total of 23 species caught in seasonal habitats, 7 contributed over 97% of total estimated numerical and biomass densities. The latter ranged from 435–580 kg ha^–1. In the 1982–83 flood-season entry to the floodplain was essentially restricted to species tolerant of oxygen-deficient waters and thereby able to penetrate the papyrus and Typha swamps of the permanent wetland. Migrants became stranded in temporary pools and subject to heavy cropping by birds.     

Cascading Ecological Responses to an In‐Stream Restoration Project in a Midwestern River

River restoration projects are increasingly common, but assessments of ecological responses and overall success of the vast majority of efforts are lacking. Information on potential positive ecological effects of restoration efforts can be used to justify further projects and refine methods. We examined responses of multiple trophic levels, aquatic insects and riparian birds, to a series of rock weirs installed in an Illinois river to stabilize the channel. We quantified adult insect emergence and performed weekly point counts of birds in spring at four weir and four non‐weir (control) sites. Emerging insect abundance was higher at control sites, but species richness and diversity were higher at weir sites. Total insect emergence production did not differ between site types, but emergence production of larger‐bodied taxa was higher at weir sites. Ordinations and analysis of similarity indicated differences in insect and bird assemblages between site types. Birds showed a positive numerical response to large‐bodied emerging insects, and total bird abundance was higher at weir sites. Clutch size and feeding rates of a focal bird species, Prothonotaria citrea (Prothonotary Warbler), did not differ between sites, but the number of hatchlings and fledglings was higher at control sites. Molothrus ater (cowbird) parasitism was higher at weir sites, likely because of increased edge habitat associated with weir construction activities. Results show positive ecological impacts of in‐stream restoration and provide justification for further efforts. However, forest disturbance associated with construction could offset some benefits to some species, and thus refinements to procedures may be necessary.     

Carbon and water cycles in tropical papyrus wetlands

Highly productive papyrus (Cyperus papyrus L.) wetlands dominate many permanently flooded areas of tropical East Africa; however, the cycling of carbon and water within these ecosystems is poorly understood. The objective of this study was to utilise Eddy Covariance (EC) techniques to measure the fluxes of carbon dioxide and water vapour between papyrus vegetation and the atmosphere in a wetland located near Jinja, Uganda on the Northern shore of Lake Victoria. Peak, midday rates of photosynthetic CO₂ net assimilation were approximately 40 μmol CO₂ m⁻² s⁻¹, while night time losses through respiration ranged between 10 and 20 μmol CO₂ m⁻² s⁻¹. Numerical integration of the flux data suggests that papyrus wetlands have the potential to sequester approximately 0.48 kg C m⁻² y⁻¹. The average daily water vapour flux from the papyrus vegetation through canopy evapotranspiration was approximately 4.75 kg H₂O m⁻² d⁻¹, which is approximately 25% higher than water loss through evaporation from open water.     

The impact of fire regimes on populations of an endangered lizard in montane south‐eastern Australia

The Blue Mountains water skink (Eulamprus leuraensis; Scincidae) is restricted to less than 40 fragmented swamp sites, all within the Blue Mountains and Newnes Plateau areas of New South Wales, Australia. Climate change is expected to increase fire frequency in the area, potentially degrading habitat quality for this endangered reptile. We quantified lizard abundances in 12 swamps using standardized surveys, and constructed a Global Information System (GIS) database to determine fire‐histories for each swamp since 1967. The abundance of Blue Mountains water skinks was negatively correlated with fire frequency, but not with time since fire. Indirect impacts of fire (mediated via shifts in vegetation density, moisture levels, prey availability and post‐fire predation) may be more important than direct effects in these cool, moist habitats. Although lizards were less common in swamps close to urban areas, and less common in frequently burnt areas, viable populations of this endangered reptile still persist even in anthropogenically disturbed swamps and in swamps that have experienced up to four fires in 20 years. Future research could usefully extend these analyses to other swamps in the locality, and explore the broader impacts of fire regimes on the distinctive flora and fauna of this threatened ecological community.     

Papyrus swamp drainage and the conservation status of their avifauna

Papyrus Cyperus papyrus swamps sustain the livelihoods of millions of people, but threats to this habitat have never been quantified formally. Birds are useful indicators of threats that cannot be measured directly. Using satellite imagery classification and habitat associated modelling, we quantify drainage and present International Union for Conservation of Nature (IUCN) Red List assessments for the eight taxa most closely associated with this habitat. We show that, between 1984–1987 and 1999–2001 the areal extent of papyrus declined by 6.7 % from 1,643 to 1,532 km². Papyrus-specialist avifauna has undergone much greater declines, due in part to fragmentation effects and in part to geographical overlap between areas of highest population densities and areas that have experienced greatest habitat loss. Our assessment does not alter the IUCN Red List status of any full species, but it improves current knowledge of the drivers of their extinction risk. Papyrus gonolek Laniarius mufumbiri should remain at least near threatened, but on the basis of population decline rather than a purported small and decreasing range size, and papyrus yellow warbler Chloropeta gracilirostris should remain vulnerable, but again due to declines rather than on the basis of previous under-estimates of population size. Other species should remain listed as least concern. However, taxonomically, likely specifically, distinct populations of papyrus yellow warbler in Zambia and Kenya are highly threatened and should be listed as endangered and critically endangered respectively. We propose several conservation priorities and discuss means of achieving these in a manner consistent with maintaining the livelihoods of people.     

Life history variation across a riverine landscape: intermediate levels of disturbance favor sexual reproduction in the ant‐dispersed herb Ranunculus ficaria

Global climate change can fundamentally alter disturbance regimes across landscapes, but little is known about how species adjust their life histories to shifts in disturbance regimes. In plants, dispersal by seeds may permit rapid re‐colonization under frequent disturbances, but often seed‐dispersing animals may be absent and local dispersal by vegetative diaspores may be a more efficient means of occupying open space in the vicinity of the plant. We tested the effect of disturbances due to inundation on the investment in seed production by the ant‐dispersed plant, Ranunculus ficaria ssp. bulbifer. During seed ripening we collected 392 plants within a landscape mosaic of 39 sites with different levels of inundation. We measured the mass of fruits and other tissues (leafs, roots, bulbils, stalks) and described plant growth form. We found that fruit numbers and masses were more variable among plants than numbers and masses of other tissues. We then standardized fruit mass against the mass of other tissues and other growth‐form parameters. Standardized fruit mass showed a highly significantly hump‐shaped relationship with the level of inundation disturbances. This pattern was consistent across 12 small‐scale transects and thus not confounded by spatial autocorrelation within landscapes. The pattern was also confirmed by analyses that simultaneously accounted for disturbance and morphological co‐variables. We thus conclude that plants invested most heavily into reproduction by seeds under intermediate levels of disturbance. Under intermediate disturbance, seeds are beneficial for rapidly re‐colonizing open space after disturbance while the seed‐dispersers are still available. The life history of mutualists such as ant‐dispersed plants and ants may thus change across a landscape, reflecting small‐scale variation in the disturbance regime.     

Banksia nectar and pollen: Dietary items affecting the abundance of the common blossom bat,Syconycteris australis,in southeastern Australia

Abstract The relationship between the abundance of the common blossom bat (Syconycteris australis), a nectar/pollen specialist, and the availability of its food sources was investigated across nine Banksia integrifolia heathland sites in coastal New South Wales, Australia. An index of bat abundance was more highly correlated with direct estimates of food density (productive inflorescences) than indirect estimates of food (Banksia trees and total inflorescences). Furthermore, standing crops of nectar-energy per hectare were better predictors of bat abundance than pollen-nitrogen per hectare. Adult male mass did not differ significantly across sites of differing food availability, suggesting that the corresponding increase in bat abundance with food reduces the amount of fat an individual can deposit. Although males and females responded similarly to food, adult numbers were more strongly correlated with food supply and they were more likely to be recaptured than juveniles. Energy requirements of blossom bats exceeded availability in four out of nine sites. Flying foxes were heard feeding in large numbers at the remaining five sites, where energy was superabundant for S. australis. In contrast to the conclusions of previous studies of megabats, these results provide the basis for a prediction that nectar-energy, rather than pollen-nitrogen, is the primary limiting factor for populations of the blossom bat.     

Avian Use of Restoration Plantings along a Creek Linking Rainforest Patches on the Atherton Tablelands, North Queensland

Rainforest fragmentation and creek degradation are major problems on the Atherton Tablelands in North Queensland. Lake Barrine and Wooroonooran National Parks are separated by a 1.5‐km length of Toohey's Creek which has been cleared except for small remnant rainforest patches. The creek has been fenced and replanted to create a corridor between the two national parks. The aim of this study was to monitor the success of the corridor in providing habitat for rainforest birds. Avian communities were monitored in the corridor and in adjacent rainforest sites using point counts over the period of reestablishment of a continuous rainforest canopy. In terms of bird community composition, within 3 years of planting, the replanted sites were on a trajectory toward the rainforest sites. Although specialist rainforest species were not yet found in the plantings, after 3 years those species which mainly inhabit rainforest but also utilize edges and riparian strips were found in similar numbers in the plantings and in the small remnant patches along the corridor. Frugivorous species were quite abundant even in the youngest plantings. Thus, replanting of rainforest trees along Toohey's Creek has begun to provide habitat for some rainforest birds. It also shows good potential to act as a corridor to connect the previously isolated patches of rainforest for these species and to develop in habitat complexity as the frugivorous birds bring in rainforest plant species from nearby areas.