The reproductive biology and early development of the African pike,Hepsetus odoe,in the Okavango Delta,Botswana

Synopsis The reproductive biology of the African pike,Hepsetus odoe, was studied over a four-year period in the Okavango Delta, Botswana. Spawning takes place between August and May in different parts of the Delta. Fish in the seasonal swamp and drainage rivers have a truncated spawning season between August and January, while those in the perennial swamp have an extended spawning season from September to May. The variation in spawning season appears to be associated with the degree of environmental perturbation brought about by the annual flood cycle. Male and female pike mature at 140 and 160 mm standard length (SL) respectively. Females predominate in the population and produce an average of 6440 eggs per season and are multiple spawners. The number of eggs per spawning averages about 2630. The eggs are deposited in a foam nest which is guarded by the parents. Newly hatched embryos suspend themselves below the nest via a cement gland and remain in the vicinity of the spawning site until they have reached a relatively advanced stage in their development. The larval period is relatively short which implies that this species undergoes nearly a direct development. Foam nests appear to be a predator avoidance mechanism as well as an adaptation to fluctuating oxygen and water levels. The initiation of the spawning season is more closely associated with water temperature than with the annual flood cycle which appears to be more important in regulating the size of the spawning stock.     

Influence of the annual flood-pulse on catch per unit effort,condition and reproduction of Clarias gariepinus from the upper Okavango Delta,Botswana

Catch per unit effort (CPUE), length, weight and maturity data for Clarias gariepinus were collected during monthly gillnet surveys in the upper Okavango Delta between 2001 and 2009 to investigate their relationship with the annual flood-pulse. CPUE, condition factor (K) and the proportion of ripe-running fish (P_RR) in the population followed a unimodal annual cycle that could be modelled using water temperature and flood-pulse hydrology. Increased CPUE during declining water levels was most likely a result of feeding migrations and aggregation behaviour. The observed increase in K during low floods in October and November preceded the increase in P_RR, which increased mainly with increasing temperature but appeared less dependent on flow. This study provided quantitative evidence that the biology of fish in the Okavango Delta is mainly dependent on the annual flood regime and, therefore, that conservation efforts should be focused on maintaining natural flow patterns in the face of climate change and potential water extraction schemes upstream.     

How landscape scale changes affect ecological processes in conservation areas: external factors influence land use by zebra (Equus burchelli) in the Okavango Delta

Most large‐bodied wildlife populations in sub‐Saharan Africa only survive in conservation areas, but are continuing to decline because external changes influence ecological processes within reserves, leading to a lack of functionality. However, failure to understand how landscape scale changes influence ecological processes limits our ability to manage protected areas. We used GPS movement data to calculate dry season home ranges for 14 zebra mares in the Okavango Delta and investigated the effects of a range of landscape characteristics (number of habitat patches, mean patch shape, mean index of juxtaposition, and interspersion) on home range size. Resource utilization functions (RUF) were calculated to investigate how specific landscape characteristics affected space use. Space use by all zebra was clustered. In the wetter (Central) parts of the Delta home range size was negatively correlated with the density of habitat patches, more complex patch shapes, low juxtaposition of habitats and an increased availability of floodplain and grassland habitats. In the drier (Peripheral) parts of the Delta, higher use by zebra was also associated with a greater availability of floodplain and grassland habitats, but a lower density of patches and simpler patch shapes. The most important landscape characteristic was not consistent between zebra within the same area of the Delta, suggesting that no single foraging strategy is substantially superior to others, and so animals using different foraging strategies may all thrive. The distribution and complexity of habitat patches are crucial in determining space use by zebra. The extent and duration of seasonal flooding is the principal process affecting habitat patch characteristics in the Okavango Delta, particularly the availability of floodplains, which are the habitat at greatest risk from climate change and anthropogenic disturbance to the Okavango's catchment basin. Understanding how the factors that determine habitat complexity may change in the future is critical to the conservation of large mammal populations. Our study shows the importance of maintaining flood levels in the Okavango Delta and how the loss of seasonal floodplains will be compounded by changes in habitat configuration, forcing zebra to change their relative space use and enlarge home ranges, leading to increased competition for key resources and population declines.     

Mapping inundation extent,frequency and duration in the Okavango Delta from 2001 to 2012

The frequency and duration of inundation in river systems are important for a variety of applications, such as water resource management, floodplain mapping and habitat restoration. The Okavango Delta, the fifth largest Ramsar site in the world, has experienced a series of large floods since 2007, following the much lower flooding of the 1990s and early 2000s. This study aimed to establish inundation frequency and duration maps of the Okavango Delta for 2001 to 2012 using MOD09Q1 and MOD11A1. An earlier methods paper testing MODIS for flood mapping in the delta showed 99.4% overall accuracy for inundation mapping of the delta, with a kappa coefficient of 80%. A method based on a time-varying threshold derived from bimodal single-band histograms was used to classify the images into inundated and non-inundated areas. The method relies on the difference in reflectance between waterbodies and dry or vegetated soil, with the former having low reflectance and the latter displaying higher reflectance values. The method takes into account seasonal changes in reflectance. Inundation maps showed a gradual decrease in the delta's maximum annual inundation extent from 2001 to 2003, followed by a gradual increase from 2004 to 2012, with maximum inundation extent during 2010–2012.     

Some observations on the geomorphological impact of hippopotamus (Hippopotamus amphibius L.) in the Okavango Delta, Botswana

The Okavango Delta is a large wetland system situated in northern Botswana. The hippopotamus forms an integral part of this dynamic ecosystem, as it exerts a catalytic effect on geomorphological change. In the distal reaches of the wetland, regular movement of hippos to feeding grounds results in the development of incised channels, which are kept clear of vegetation and act as nodes for swamp expansion. Hippos maintain pathways in backswamp areas, which lead to the development of new channel systems during channel avulsion. They create breaches in the vegetation levees which flank channels in the permanent swamps, causing diversion of water and sediment to backswamp areas. Their paths often lead to lakes in the permanent swamps. During channel avulsion, diversion of a channel into lakes may occur via these paths, which can result in lake closure.     

Potential environmental impacts associated with the proposed abstraction of water from the Okavango River in Namibia

The Namibian Department of Water Affairs has in the past faced considerable pressure to relieve the water shortages caused by recent droughts. One of the options considered in 1996, following poor runoff during the 1994/95 and 1995/96 seasons, was a proposal to abstract some 17Mm³ of water per year from the Okavango River at Rundu, and transfer this via a 260km long pipeline to the head of the Eastern National Water Carrier at the town of Grootfontein. Part of the overall evaluation of this scheme included an assessment of the potential environmental impacts that could arise. An initial environmental evaluation was conducted from a point approximately 40 kilometres upstream of Rundu in Namibia, to the distal end of the Okavango Delta at Maun in Botswana.

Hydrological studies showed that the proposed abstraction represented a reduction of approximately 0.32% in the mean annual flow of the Okavango River at Rundu. The abstraction represents 0.17% of the mean annual flow at Mukwe, downstream of the Cuito River confluence. The adverse effects of the proposed water abstraction scheme would be extremely small along the Okavango River in Namibia, whilst outflows from the lower end of the Okavango Delta to the Thamalakane River would be reduced by some 1.44Mm³/year (11%). Additional studies showed that these effects could be reduced by some 10–13% if abstraction was confined to the falling limb of the hydrograph.

Hydrological simulations have shown that the maximum likely loss of inundated area in the Okavango Delta would total approximately 7km² out of some 8 000km². This potential loss in inundated area would be concentrated in the lower reaches of the seasonal swamps and seasonally inundated grasslands, specifically in the lower reaches of the Boro, Gomoti, Santantadibe and Thaoge channels. However, these effects would most likely be expressed as a shoreline effect, with the loss in area spread out over the shoreline and periphery of islands and would not be restricted to a single specific area. This anticipated loss in inundated area is unlikely to have measurable impacts on environmental components.

Overall, the study found no 'fatal flaws' which would prevent the water abstraction scheme from proceeding and the anticipated effects on the Okavango system are more likely to be seen in the Okavango Delta, rather than along the Okavango River. The anticipated ecological implications of the scheme were small in spatial extent and are unlikely to be perceptible against the natural year-to-year variability in inundation of the Okavango Delta or outflows to the Thamalakane River. However, the public perceptions of the proposed water transfer project were strongly negative and appeared to be at least in part due to the very low water levels in the Okavango River and Okavango Delta during the past three years and during the study period. These low water levels were amongst the lowest on record and it is likely that the public would attribute any adverse effect recorded in the future to the abstraction scheme, whether this were true or not. These negative perceptions of the desirability and acceptability of the proposed scheme were strongly linked to potential adverse affects on the tourism industry along the Okavango River and in the Okavango Delta, with possible adverse economic effects on local residents.     

FISH AND FISHERIES OF WETLANDS

SUMMARY

The three major wetland types have characteristic fish communities. River-associated wetlands harbour a rich diversity of fishes which can either survive habitat desiccation during the dry-down, or migrate to and fro with the flood. Endorheic wetlands have a lower diversity of species which are typically ‘r’-selected relative to mainstream riverine or lacustrine forms. Marine wetlands have a variety of peripheral or marine forms, but are not discussed in this account. The dynamics of wetland fish communities are determined by periodically changing abiotic factors, especially water temperature and water level, and biotic factors, especially food availability. Water level fluctuations have several important functions and result in pulses of nutrient input and fish abundance. Wetland fish stocks can usually be sustained as long as the pristine flood regime is retained, but disruption of the flooding pattern interferes with fish breeding and nutrient flow. Fishing yields may be directly correlated with the flood history during the previous two years. Wetland fishes play an important role by converting the resource at the base of the food chain, i.e. living plants, detritus or epiphytes, into food for higher trophic levels. The need to identify and conserve key food supplies is emphasized. Wetland fisheries are relatively poorly developed in southern Africa with notable exceptions, e.g. Pongolo floodplain, Okavango swamp, Lake Liambezi, Elephant Marsh on the Shire River, but even in these systems the resource is probably underutilized.

The need to consider the socio-economic implications of upgrading a subsistence fishery are emphasized, and further study on the effect of water level fluctuations on major trophic pathways and on multispecies fisheries is encouraged. Because of the urgent need to manage wetland fisheries, various short-cut methods to obtain first-order estimates of fish production, yields and growth rates should be used until more data are available. The need to manage a floodplain holistically for both the aquatic and terrestrial phases is stressed. In addition, inocula (re-colonizing life history stages) should be conserved to accelerate the recovery of endorheic wetland fish stocks after a drought. As wetlands are of fundamental importance to lake or river metabolism and provide vital breeding and feeding areas for fishes, they should be given the highest conservation priority. Cost-benefit analyses should therefore be performed before any large scale manipulations of wetlands are performed.     

The reproductive and feeding biology of Schilbe intermedius Rüppell in the Okavango Delta,Botswana

The reproductive and feeding biology of 314 Schilbe intermedius Rüppell 1832 collected in the Okavango Delta, Botswana, between April 1985 and February 1986 during low, intermediate and high water level were investigated. S. intermedius matures at 130 mm standard length and is a monocyclic spawner. Females predominate in the population and have an average fecundity of 27107 eggs. In the northern riverine floodplain of the Delta fish spawn in mid-summer during the peak flood. In the southern drainage rivers fish spawn in spring, which is two to three months after the arrival of the winter flood waters. S. intermedius is an opportunistic predator with a diet consisting largely of fish 41%, aquatic larvae 25%, terrestrial insects 14%, aquatic insects 7% and crustacea 5%. Juvenile S. intermedius feed predominantly on insect larvae, while adults are largely piscivorous.     

Do steady state assemblages occur in shallow lentic environments from wetlands?

The Okavango Delta has an annual flood cycle that spreads slowly from the northwestern`panhandle' into a widening delta to the southeast. Rotifers were sampled from areas where the flood was active, and from other areas not yet reached by the flood. In still waters, the samples were dominated by euplanktonic rotifers, particularly of the genera Brachionus, Keratella and Hexarthra, with about 20 species per 1000 individuals at a station. Where the water was moving the samples were dominated by periphytic rotifers, with over 100 species per 1000 individuals at a single station. The periphytic species were dominated by the genus Lecane,with over 45 species, followed by Lepadellawith 24 species, and Trichocercawith 23 species. Comparison of the Okavango with six other tropical localities reveals fairly consistent ratios of the numbers of species in the periphytic genera Lecane, Lepadellaand Trichocerca,indicating that there may be assembly rules for communities of tropical periphytic rotifers.     

Two genotypic groups of morphologically similar fish trypanosomes from the Okavango Delta, Botswana

Blood smears and blood lysate samples from freshwater fishes captured in the Okavango Delta, Botswana, were examined to determine whether their trypanosomes were all Trypanosoma mukasai, a species of supposed broad host specificity and widespread existence across Africa. Trypanosomes and/or babesiosomes occurred in 20/32 blood smears, and morphometric analysis of trypanosomes from 13/32 smears showed features suggestive of T. mukasai, including nuclear indices consistently >1. In 16/32 blood lysate samples from which DNA was extracted, trypanosome DNA was detected in 12/16 by PCR (polymerase chain reaction), using trypanosome-specific ssu rRNA gene primers. Two samples positive for trypanosomes in blood smears yielded no amplifiable trypanosome DNA, but 4 samples with no detectable infection in blood smears were positive for trypanosome DNA, suggesting an overall trypanosome prevalence rate of 17/32 (53%) among fishes and demonstrating the value of PCR in trypanosome recognition. Cloning and sequencing of the 12 amplified fragments revealed 2 genotypic groups among these fish trypanosomes. Group 1 trypanosomes were from cichlids and 3 families of catfishes, Group 2 from 2 types of catfishes. Sequence comparison showed that the consensus Group 1 sequence was most similar to that of Trypanosoma cobitis, representing European fish trypanosomes of the carassii type, while the consensus Group 2 sequence showed similarity with a trypanosome sequence from another African catfish, Clarias angolensis. It was concluded that the identification of T. mukasai remains a problem, but at least 2 genotypic groups of trypanosomes occur in Okavango Delta fishes, and catfishes in this region appear to contain both types.     

Guidelines for RNA-seq projects: applications and opportunities in non-model decapod crustacean species

Understanding and predicting vegetation patterns in floodplains are essential for conservation and/or restoration of river floodplains subject to hydrological alterations. We propose a conceptual hydroecological model to explain the disturbance mechanisms driving species diversity across large river floodplains. These ecosystems harbor a unique set of flood-tolerant species different from the surrounding upland vegetation. In elevation gradients across pristine floodplains, the greater the flooding, the fewer the number of plant species. As terrain elevation increases, flood depth and duration decrease and it is more likely that species composition is influenced by external natural or human-driven disturbances. The spatial interaction between the natural flood regime and upland factors creates patterns of disturbance gradients that influence how floodplain vegetation establishes. In regions where upland conditions are subject to strong external disturbances, species diversity peaks at intermediate stages along the disturbance gradient. We demonstrate this concept with observations from the Central Amazon and Pantanal in Brazil, the Mekong’s Tonle Sap in Cambodia, and the Okavango Delta in Botswana. We discuss how this model could be further elaborated and validated to inform management of large river basins under the impact of upstream-induced flood pulse alterations.     

The relevance of fire frequency for the floodplain vegetation of the Okavango Delta, Botswana

The study investigates the relevance of fire frequency for the floodplain vegetation of the Okavango Delta in Botswana, considering species composition, species richness, vegetation structure and tree density. Based on flood and fire frequency data derived from a series of satellite images, active and drying floodplains were separated and study plots were selected with fire frequencies up to ten burns between 1989 and 2003. Both for drying and active floodplains, no typical species assemblage could be associated with fire frequency. On drying floodplains, fire frequency showed no effect on species composition, but rather on vegetation structure. While small woody species showed higher cover values on high fire frequency, large trees showed significantly lower cover values on high fire frequency. For active floodplains, a significant response to fire frequency, both positively and negatively could be determined for specific species. But as almost none of these species appear at low or high frequency only, no differences in species composition could be associated with fire frequency. Rather flood frequency and specific annual flood cycles could be described as the determining factors for the vegetation on active floodplains.     

River-wetland interaction and carbon cycling in a semi-arid riverine system: the Okavango Delta, Botswana

The Okavango River, in semi-arid northwestern Botswana, flows for over 400 km in a pristine wetland developed on a large (>22,000 km²) alluvial fan (Okavango Delta). An annual flood pulse inundates the floodplains of the wetlands and travels across the Delta in 4–6 months. In this study, we assess the effects of long hydraulic residence time, variable hydrologic interaction between river–floodplain–wetland and evapotranspiration on carbon cycling. We measured dissolved inorganic carbon (DIC) concentrations and stable carbon isotopes of DIC (δ¹³C_DIC) from river water when the Delta was not flooded (low water) and during flooding (high water). During low water, the average DIC concentration was 31 % higher and the δ¹³C_DIC 2.1 ‰ more enriched compared to high water. In the lower Delta with seasonally flooded wetlands, the average DIC concentration increased by 70 % during low water and by 331 % during high water compared to the Panhandle with permanently flooded wetlands. The increasing DIC concentration downriver is mostly due to evapoconcentration from transpiration and evaporation with increased transit time. The average δ¹³C_DIC between low and high water decreased by 3.7 ‰ in the permanently flooded reaches compared to an increase of 1.6 ‰ in the seasonally flooded reaches. The lower δ¹³C_DIC during high water in the permanently flooded reaches suggest that DIC influx from the floodplain-wetland affects river’s DIC cycling. In contrast, higher river channel elevations relative to the wetlands along seasonal flooded reaches limit hydrologic interaction and DIC cycling occurs mostly by water column processes and river-atmospheric exchange. We conclude that river-wetlands interaction and evapoconcentration are important factors controlling carbon cycling in the Okavango Delta.     

长三角城市群城市洪涝韧性与生态系统服务的耦合协调关系

厘清城市洪涝韧性与生态系统服务之间耦合协调关系,可为城市防洪减灾和生态文明建设提供重要决策参考。综合运用基于麻雀算法的投影寻踪模型、InVEST模型和耦合协调度模型,在分析2000-2020年长三角城市群的城市洪涝韧性与水源涵养、水质净化、土壤保持和气候调节四种生态系统服务时空格局基础上,尝试探索两者耦合协调关系及其时空演变特征。研究发现:(1)长三角城市群的城市洪涝韧性水平呈现"N"型增加趋势,并呈现"上海>江苏>浙江>安徽"的空间格局,表明经济水平越高的城市展现出更强的洪涝韧性,经济是影响城市洪涝韧性波动的主要因素,而自然韧性成为城市洪涝韧性提升的关键短板;(2)生态系统服务存在显著的空间异质性,高植被覆盖的南部地区提供了更高的生态系统服务,从时间维度看其空间分布是稳定的,水源涵养和水质净化服务总体呈向好趋势,土壤保持服务整体呈现倒"N"型增加,气候调节服务呈现微弱的下降趋势;(3)城市洪涝韧性与生态系统服务的耦合协调度在研究期内较为稳定,且与生态系统服务的时空变化趋于一致,呈现"南高北低、由西南向东北逐渐减弱"的趋势,生态系统服务较高的城市,表现出更好的耦合协调性,且随着生态系统服务的增加而改善。因此,长三角城市群有必要从提高生态系统服务功能的角度出发,因地制宜、分类施策,促进城市洪涝韧性与生态系统服务的协调发展。     

Freshwater snails of East Caprivi and the lower Okavango River Basin in Namibia and Botswana

A systematic account is given of the aquatic gastropod fauna of the lower Okavango River in Namibia and Botswana, and of the East Caprivi area in Namibia, based on collections made mostly in 1983–86 from about 100 different sites. A total of 20 living species are reported, 9 of them for the first time from this area: Bellamya monardi, Lobogenes michaelis, Cleopatra elata, Afrogyrus coretus, Segmentorbis angustus, S. kanisaensis, Bulinus scalaris, B. depressus and B. tropicus. All are found outside this area and most are widely distributed Afrotropical species. Some do not occur farther south than the Okavango Delta, while others reach a south-western limit here but occur at greater latitude in the eastern tropical corridor on the coast of Natal. Bulinus globosus and Biomphalaria pfeifferi occur throughout the study area and are intermediate hosts for schistosome parasites of man and livestock.     

Algal richness and life‐history strategies are influenced by hydrology and phosphorus in two major subtropical wetlands

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An Elephant in the Room: Okavango Safari Hunting as Ecotourism?

Safari hunters’ acute awareness of the widely held negative perceptions of their practice has led to their development of strong justifications and defensive assertions in favour of hunting. Far from being a primarily destructive practice, they claim that safari hunting in the Okavango Delta, Botswana, can be seen as an exemplary form of ecotourism, which benefits local communities, facilitates environmental conservation and provides the ultimate nature experience for participants. While research supports their claims to an extent, the ethical quandaries evinced by hunters themselves, the complex dialectic between local and global controls, and the elite, racialised and gendered nature of hunting speaks to a more complex and conflicted situation.     

Physiological refugia: swamps,hypoxia tolerance and maintenance of fish diversity in the Lake Victoria region

In Lake Nabugabo, Uganda, a satellite of Lake Victoria, approximately 50% of the indigenous fishes disappeared from the open waters subsequent to the establishment of the introduced predatory Nile perch, Lates niloticus. This pattern is similar to the faunal loss experienced in the much larger Lake Victoria. Several of these species persisted in wetland refugia (e.g. ecotonal wetlands, swamp lagoons); however, deep swamp refugia (habitats lying well within the dense interior of fringing wetlands), are available only to a subset of the basin fauna with extreme tolerance to hypoxia. Although air-breathers are common in deep swamp refugia; we also documented a surprisingly high richness and abundance of non-air-breathing fishes. We describe several mechanisms that may facilitate survival in deep swamp refugia including high hemoglobin concentration, high hematocrit, large gill surface area and a low critical oxygen tension (P(c)). In addition, swamp-dwelling fishes showed lower PO(2) thresholds for onset of aquatic surface respiration than the lake-dwelling fishes. This suggests higher tolerance to hypoxia in the swamp fishes because they are able to withstand a lower oxygen tension before approaching the surface. We suggest that physiological refugia may be important in modulating the impact of Nile perch and indigenous fishes in the Lake Nabugabo region; this highlights the need to evaluate relative tolerance of introduced predators and indigenous prey to environmental stressors.     

Consistent trophic patterns among fishes in lagoon and channel habitats of a tropical floodplain river: Evidence from stable isotopes

The relationship between food web dynamics and hydrological connectivity in rivers should be strongly influenced by annual flood pulses that affect primary production dynamics and movement of organic matter and consumer taxa. We sampled basal production sources and fishes from connected lagoons and the main channel of a low-gradient, floodplain river within the Orinoco River Basin in Venezuela. Stable isotope analysis was used to model the contribution of four basal production sources to fishes, and to examine patterns of mean trophic position during the falling-water period of the annual flood cycle. IsoSource, a multi-source mixing model, indicated that proportional contributions from production sources to fish assemblages were similar in lagoons and the main channel. Although distributions differed, the means for trophic positions of fish assemblages as well as individual species were similar between the two habitats. These findings contradict recent food web studies conducted in temperate floodplain rivers that described significant differences in trophic positions of fishes from slackwater and floodplain versus main channel habitats. Low between-habitat trophic variation in this tropical river probably results from an extended annual flood pulse (ca. 5 mo.) that allows mixing of sestonic and allochthonous basal production sources and extensive lateral movements of fishes throughout the riverscape.     

Pack-hunting in two species of catfish, Clavias gariepinus and C. ngamensis, in the Okavango Delta, Botswana

During the annual drawdown in the Okavango Delta, Botswana, large shoals of catfish migrate upstream in the main river channels. The examination of samples of the sharptooth catfish Clarias gariepinus and blunttooth catfish C. ngamensis revealed that they were pack-hunting and selectively preying on two species of mormyrids, the bulldog Marcusenius macrolepidotus and the churchill Petrocephalus catostoma . The predation on mormyrids may be related to prey size and abundance. In addition, catfish may sense the electrical discharge given off by the mormyrids when disturbed. The catfish feed intensively for a few months and build up body reserves and reproductive condition prior to spawning on shallow floodplains with the onset of the annual floods from Angola.