Jude M. Mathooko

A global experiment suggests climate warming will not accelerate litter decomposition in streams but might reduce carbon sequestration

The decomposition of plant litter is one of the most important ecosystem processes in the biosphere and is particularly sensitive to climate warming. Aquatic ecosystems are well suited to studying warming effects on decomposition because the otherwise confounding influence of moisture is constant. By using a latitudinal temperature gradient in an unprecedented global experiment in streams, we found that climate warming will likely hasten microbial litter decomposition and produce an equivalent decline in detritivore-mediated decomposition rates. As a result, overall decomposition rates should remain unchanged. Nevertheless, the process would be profoundly altered, because the shift in importance from detritivores to microbes in warm climates would likely increase CO(2) production and decrease the generation and sequestration of recalcitrant organic particles. In view of recent estimates showing that inland waters are a significant component of the global carbon cycle, this implies consequences for global biogeochemistry and a possible positive climate feedback.

Macroinvertebrates: Composition, Life Histories and Production

Publisher Summary This chapter describes the composition of the macroinvertebrate fauna of tropical streams, especially the insects, with the aim of uncovering the patterns in diversity and richness that they exhibit. It reveals that insects and other tropical macroinvertebrates are sometimes affected by temperature, but that flow regime and habitat stability have a primary influence on life-history parameters and population dynamics. Decapods make an important contribution to overall production in some tropical streams, and may offset low production by insects. “Macroinvertebrates” are those invertebrates exceeding 0.5 mm body size, or large enough to be seen by the naked eye, and comprise mostly insects as well as decapod crustaceans, mollusks, leeches, oligochaetes, and planarians. The none -insect groups are dominated by turbellarian Planariidae, mollusks, and crustaceans. It shows that most studies of the life-history attributes of macroinvertebrates in tropical streams concern the temporal components of voltinism and phenology and most have concerned insects. Macroinvertebrate secondary production estimates integrate information on population dynamics and energy flow that are of considerable ecological importance. One obvious gap in studies of macroinvertebrate lifehistories is uncertainty about fate and behavior of adult aquatic insects in the tropics. It highlights that the amounts of research devoted to macroinvertebrates in tropical Africa is much less than elsewhere in the tropics, yet the threats that streams in this region face are no less than those in the rest of the tropics.

Organic Matter Processing in Tropical Streams

Organic matter derived from many sources provides a basis for stream food webs. In terms of weight, leaves from the surrounding land constitute the largest allochthonous source of energy for stream consumers, but other items, including fruits, flowers, wood and twigs, and terrestrial insects, are also important. Timing of allochthonous inputs can vary markedly due to the phenology of the riparian vegetation, retention mechanisms in the aquatic-terrestrial transition zone, and local climate (especially the incidence of high-rainfall events), but seasonality of litter inputs is different, and often much less marked, than is typical of streams in temperate latitudes. As in such streams, litter decomposition rates depend on the interaction of physical factors (flow, temperature), water chemistry (dissolved nutrients), and biological agents (micro-organisms and detritivores – especially shredding invertebrates). Because vascular plant biodiversity in the tropics is high, varied leaf characteristics (hardness, phenolic content, and other aspects of leaf chemistry) contribute to great variability in breakdown rate: fast-decomposing leaves persist for a few days only, whereas highly recalcitrant species take well over a year to decompose. In all the above cases, the decomposition process includes an initial rapid leaching phase when water-soluble compounds are lost, followed by colonization by micro-organisms (fungi and bacteria), and subsequent mechanical breakdown of the leaf structure by invertebrate shredder and hydraulic forces. Undecomposed leaves are sometimes exported downstream during flood events, and thence deposited in water-logged riparian zones or, in some cases, forming dense accumulations of peat that are important as carbon sinks and as habitat for specialized biota. Recent research indicates that the role of invertebrate shredders in processing organic matter in tropical streams is less than in temperate latitudes, and there may be a higher proportion of material that is recalcitrant and/or exported from streams (or stored as peat) before it is decomposed completely. Autochthonous energy sources may be particularly important to consumers in tropical streams, and there is some evidence of a lesser reliance on allochthonous organic matter than in temperate streams.

Disturbance of a Kenya Rift Valley stream by the daily activities of local people and their livestock

Human and domestic animal activities and visits to an impacted site in the Njoro River, Kenya, were recorded from 1994 to 1995. The activities of people in the humid and wet zones of the stream included linen washing, water abstraction, excretion, bathing and swimming. Human and animal visits and activities along the Njoro River were on a daily basis and were patterned according to the time of the day, weather and seasons. Women formed the first group to visit the stream at dawn, followed by men and lastly, children. The diurnal pattern of visits was bimodal, with major peaks between 0600 and 1100 h and from 1600 h until dusk. The intensity of all major activities peaked at around midday and donkeys and cattle formed the largest proportion of the domestic animals that visited the stream. Much of the small-scale water abstraction occurred between 0700 and 1100 h, coinciding with the time when most people visited the stream. Water abstraction was most intense during the dry season. A mean discharge of 1.7 litres per second was measured whilst 0.3 litre of water per second was abstracted at the impacted site, implying that about 20% of the flow volume of the Njoro River was abstracted at a single site in one day.The effects of the human and animal activities on the structure of macrozoobenthos in the wet zone of the Njoro River were studied on the ‘impacted site’ in relation to an ‘upstream reference site’ and a ‘downstream reference site’. Oligochaetes and chironomids dominated the fauna in all three sites. The impacted site had low patchiness and mean crowding, with the taxa distribution tending toward a randomly dispersed spatial pattern. The mean turnover (±SD) of the macroinvertebrates was 47±18, 48±26 and 36±22 in the impacted, downstream and upstream sites, respectively. The trampling of the streambed by humans and livestock could, therefore, alter the benthos structure through redistribution and reduction of faunal patchiness. The cumulative effect of small-scale, but widespread and frequent disturbances might have large impacts on whole river systems. This study demonstrated that, in the tropics, quantification of the daily activities of people and domestic animals is important for future management of the Njoro River and consequently, Lake Nakuru.

Leaf litter processing rates in a Kenyan highland stream, the Njoro River

Mass loss of leaf litter from five common riparian trees in Kenya was determined using mesh bags over a 70-day period. The study was carried out in a river which previous research has shown to be apparently devoid of invertebrate shredders. There was a five fold difference between the most rapid and slowest decay rates, but all were within the fast range identified from temperate zone trees, demonstrating high rates of microbial activity. We conclude, therefore, that the low biomass of invertebrate shredders in Kenyan highland streams cannot be explained by poor quality of the leaf litter that enters them.

Composition and seasonality of benthic invertebrates, and drift in the Naro Moru River, Kenya

A survey was carried out to establish the nature and composition of the benthos along the Naro Moru, a tropical river in central Kenya using artificial substrate baskets, from November 1986 to October 1987. A clear longitudinal zonation existed for Diptera and Ephemeroptera which were the major benthic taxa. Maximum colonization took place after ten days of exposure.Seasonal variations in abundance were also observed. All taxa collected from the bottom samples were also collected in the drift samples, but the percentage composition of the benthos showed variations with that of the drift. Simulium sp. dominated the benthos whilst Baetis spp. dominated the drift. There was a positive correlation between drift rate and benthic fauna density.

Distribution and abundance of freshwater crabs ( Potamonautes spp.) in rivers draining Mt Kenya, East Africa

Little is known about population densities of freshwater crabs, or their ecological importance, in African rivers. This study aimed to quantify crab abundance in rivers draining Mt Kenya. Invertebrates were sampled from 21 sites on 19 rivers. Sample sites were divided into: open sites in agricultural land from which trees were mainly absent; shaded sites, in agricultural land, with cultivated catchments but heavy shading by riparian trees; and forest sites, still dominated by natural vegetation. Crabs, mainly Potamonautes odhneri, were recorded from 14 of the 21 sites, including all forest sites, and were significantly more abundant in forest sites than in either type of agricultural site. However, there was no difference in biomass among habitat types, because individual crabs were significantly smaller on average in forest sites. This was due to the large numbers of small juvenile crabs recorded in forest sites, whereas small juveniles were almost absent from all agricultural sites. Although numerically unimportant relative to other macroinvertebrates, crabs accounted for at least 70 % of total macroinverte-brate biomass from forest and shaded agricultural sites, and averaged around 40 % in open agricultural sites. It is possible that crab reproduction occurs mainly or exclusively in forested areas, which would therefore act as a recruitment source for populations farther downstream in agricultural areas.

Leaf-litter breakdown in tropical streams: is variability the norm?

Many forested headwater streams are heterotrophic ecosystems in which allochthonous inputs of plant litter are a major source of energy. Leaves of riparian vegetation entering the stream are broken down by a combination of biotic and abiotic processes and, in most temperate and boreal streams, provide food and habitat for dense populations of detritivorous invertebrates. However, tropical streams in different parts of the world show substantial variability in the number and diversity of leaf-shredding detritivores (hereafter detritivores). We used data obtained with standardized methods from multiple streams in Africa, the Americas, Asia, and Australia to test the hypothesis that this variability would lead to differences in the relative role of detritivores and microorganisms in the breakdown process. We also tested the hypotheses that variability in litter breakdown rates changes with litter type (native litter mixtures vs nonnative alder [Alnus glutinosa]) and is higher across regions within than outside the tropics. We found that litter breakdown rates were highly variable across sites, with no consistent pattern within geographic areas, although litter consumption by detritivores was negligible at several sites, all in America. Geographic patterns of litter breakdown also varied between litter types, with higher breakdown rates for alder than for native litter in most but not all regions. When litter breakdown rates at the tropical sites were compared to previously reported values from temperate and boreal regions, we found that differences in variability between tropical and temperate sites were inconsistent, with great differences among studies. Further global-scale studies will be needed to assess the extent to which latitudinal changes in the diversity and composition of microbial and detritivore assemblages contribute to variability in litter breakdown rates.

Decomposition of leaf litter of Dombeya goetzenii in the Njoro River, Kenya

Decomposition of the leaves of Dombeya goetzenii (K. Schum) in the Njoro River is described and analysed. The loss of the ash-free dry mass was rapid during the first 14 d of exposure in the wet and humid zones. The leaves in the litter bags in the humid and wet zones were processed at a rate (±SD) of 0.005±0.001 d-1 and 0.021±0.001 d-1, respectively. The processing rates of the leaves in the wet zone differed significantly from those observed in the humid zone (t-value, p<0.05). The interchanged litter bags (i.e. from wet to humid zones vice versa) showed that the processing rates of the leaves in the litter bags interchanged from the wet zone to the humid zone was about 200 times lower than that of the leaves retained in the wet zone throughout; fourfold higher in the leaves in the litter bags which were transferred to the wet zone than in those leaves of the litter bags which were retained in the humid zone throughout the experiment. It took approximately 38 months for 90% of the leaf dry mass to be processed in the humid zone whilst it took 4 months for a similar percentage to be processed in the wet zone. It is concluded that the immersion and emersion of leaf litter, which may occur in the wet and humid zones, respectively, are important aspects of the decomposition process which may influence the quantity of nutrients in stream ecosystems.

Leaf litter transport and retention in a tropical Rift Valley stream: an experimental approach

The dynamics of leaf litter transport and retention in two contrasting reaches in the Njoro River, Kenya, is reported, analysed and discussed. The study focussed on two aspects: experimental leaf litter transport and retention in the Maryjoy and Turkana reaches which differ in structural heterogeneity and natural leaf litter transport in the two reaches. Leaves of Dombeya goetzenii, Syzygium cordatum and Grewia forbesii were dyed differently with ‘Herbol®’ dyes and used to elucidate the dynamics of leaf transport and retention. Experimentally, about 40% of each of the three leaf species released into the water column travelled beyond the experimental 50-m mark in the Maryjoy reach (devoid of major retention structures) and about 95% of the leaves were retained within the Turkana reach (with major retention structures) and never reached the 50-m mark. Based on the calculated average leaf travel distance (1/k), the leaves of D. goetzenii could have drifted 10–113 m in the Turkana reach and between 60 and 127 m in the Maryjoy reach. The transport trends of the three leaf species were reach- but not leaf species-specific. The efficiency of trapping the drifting leaves differed from one retention structure to another in both reaches. Rock outcrops and debris dams were the main descriptors of the stream heterogeneity in the Turkana reach and retained about 55% and 20% of the experimental leaves, respectively. The leaves of S. cordatum dominated the leaf litter transported naturally within the two reaches, contributing 214.4±18.7 g d−1 to 274.1±36.8 g d−1 (± standard error) of the leaf litter biomass in the Turkana reach. It was estimated that 640.9 kg of leaf litter are exported to Lake Nakuru by the Njoro River annually.