Olaf L. F. Weyl

Advancing impact prediction and hypothesis testing in invasion ecology using a comparative functional response approach

Invasion ecology urgently requires predictive methodologies that can forecast the ecological impacts of existing, emerging and potential invasive species. We argue that many ecologically damaging invaders are characterised by their more efficient use of resources. Consequently, comparison of the classical ‘functional response’ (relationship between resource use and availability) between invasive and trophically analogous native species may allow prediction of invader ecological impact. We review the utility of species trait comparisons and the history and context of the use of functional responses in invasion ecology, then present our framework for the use of comparative functional responses. We show that functional response analyses, by describing the resource use of species over a range of resource availabilities, avoids many pitfalls of ‘snapshot’ assessments of resource use. Our framework demonstrates how comparisons of invader and native functional responses, within and between Type II and III functional responses, allow testing of the likely population-level outcomes of invasions for affected species. Furthermore, we describe how recent studies support the predictive capacity of this method; for example, the invasive ‘bloody red shrimp’ Hemimysis anomala shows higher Type II functional responses than native mysids and this corroborates, and could have predicted, actual invader impacts in the field. The comparative functional response method can also be used to examine differences in the impact of two or more invaders, two or more populations of the same invader, and the abiotic (e.g. temperature) and biotic (e.g. parasitism) context-dependencies of invader impacts. Our framework may also address the previous lack of rigour in testing major hypotheses in invasion ecology, such as the ‘enemy release’ and ‘biotic resistance’ hypotheses, as our approach explicitly considers demographic consequences for impacted resources, such as native and invasive prey species. We also identify potential challenges in the application of comparative functional responses in invasion ecology. These include incorporation of numerical responses, multiple predator effects and trait-mediated indirect interactions, replacement versus non-replacement study designs and the inclusion of functional responses in risk assessment frameworks. In future, the generation of sufficient case studies for a meta-analysis could test the overall hypothesis that comparative functional responses can indeed predict invasive species impacts.

The effect of stocking density on growth, survival and agonistic behaviour of African catfish

A high rate of sibling cannibalism is one of the principal obstacles in the rearing of larvae and juveniles of the African catfish Clarias gariepinus. This paper examines the underlying behavioural components of agonistic behaviour under culture conditions. Ten day old catfish larvae were stocked at 30 larvae l−1 in three tank designs with different surface areas and equal volumes. Stocking densities were 1.2, 0.6 and 0.3 fish cm−2 bottom surface. Growth did not differ between treatments. The highest mortality over 30 days was recorded at the medium density (0.6 larvae l−1). Fish stocked at this density showed the highest rate of aggression, while there was no difference in aggression between the highest and the lowest stocking densities. The results indicate that stocking density should be at least as high as 1.2 larvae cm−2 bottom surface area to obtain high production at best survival rates.

Existing and emerging high impact invasive species are characterized by higher functional responses than natives

Predicting ecological impacts of invasive species and identifying potentially damaging future invaders are research priorities. Since damage by invaders is characterized by their depletion of resources, comparisons of the ‘functional response’ (FR; resource uptake rate as a function of resource density) of invaders and natives might predict invader impact. We tested this by comparing FRs of the ecologically damaging ‘worlds worst’ invasive fish, the largemouth bass (Micropterus salmoides), with a native equivalent, the Cape kurper (Sandelia capensis), and an emerging invader, the sharptooth catfish (Clarias gariepinus), with the native river goby (Glossogobius callidus), in South Africa, a global invasion hotspot. Using tadpoles (Hyperolius marmoratus) as prey, we found that the invaders consumed significantly more than natives. Attack rates at low prey densities within invader/native comparisons reflected similarities in predatory strategies; however, both invasive species displayed significantly higher Type II FRs than the native comparators. This was driven by significantly lower prey handling times by invaders, resulting in significantly higher maximum feeding rates. The higher FRs of these invaders are thus congruent with, and can predict, their impacts on native communities. Comparative FRs may be a rapid and reliable method for predicting ecological impacts of emerging and future invasive species.

On the Life History of a Cyprinid Fish, Labeo cylindricus

Aspects of the life history of Labeo cylindricus in Lake Chicamba, a man made hydroelectric dam in central Mozambique were investigated. L. cylindricus was found to be a potadrometic, synchronous spawner concentrating in river mouths prior to migrating up flowing rivers in January to spawn. Length-at-(50%)-maturity was attained at a fork length (FL) of 96 mm for males and 98 mm for females, both within their first year of life. The adult sex ratio was found to be female dominated at 1 male:1.63 females. Scales were used to age the species, with two growth checks deposited annually. Growth was rapid and was best described by the von Bertalanffy growth model as Lt=224.2(1−e-0.66(r+0.03)) mm FL. A maximum age of 4 years was observed for L. cylindricus with the largest male and female measuring 210 mm FL and 235 mm FL, respectively. The rate of natural mortality was estimated to be extremely high at 1.93 year-1 using length-converted catch curve analysis and is possibly a response to heavy predation by the introduced piscivore Micropterus salmoides.

Threatened Endemic Fishes in South Africa's Cape Floristic Region: A New Beginning for the Rondegat River

ABSTRACTNonnative fishes threaten native fish communities in many rivers of the world. In South Africas Cape Floristic Region, predation by nonnative fishes has severely impacted native fish populations and more than half of the 17 endemic fish species are endangered. To preserve the unique endemic fish fauna, removal of nonnative fish from conservation areas is a priority in this region. In February 2012, South Africas first nonnative fish eradication using rotenone took place in the Rondegat River, a small headwater stream that had been invaded by Smallmouth Bass (Micropterus dolomieu). The successful treatment culminated from a decade-long process that was facilitated through collaboration among a South African nature conservation authority (CapeNature), the South African Institute for Aquatic Biodiversity, and the American Fisheries Society Fish Management Chemicals Subcommittee. The successful removal of alien fish and almost instantaneous increase in biodiversity is anticipated to encourage more e...

A successful population of largemouth bass, Micropterus salmoides, in a subtropical lake in Mozambique

Sectioned otoliths were used for age and growth determination of the alien Micropterus salmoides from Lake Chicamba, a hydroelectric dam in central Mozambique. Marginal zone analysis showed that annulus formation occurred during winter in August and September. The maximum estimated age was 5 years. Growth was described by the 3 parameter von Bertalanffy model as Lt=465.51 (1 − e-1.175(t+0.009)) mm FL. Juvenile growth rate was 23.1 mm per month during their first 7 months of life. The age at 100% maturity was 0.9 years, and spawning occurred during August and September. The juvenile fish fed sequentially on the most abundant prey items. Spawning occured before any of the indigenous fish species and so, there is little inter-specific competition. Conspecific juveniles formed the most important constituent in the adult M. salmoides diet from October to December when alternative prey was limited. The total annual mortality rate (Z) for the species in Lake Chicamba was 1.27 yr-1, the mean empirical estimate of natural mortality (M) was 0.73 yr-1, and fishing mortality (F) was calculated at 0.54 yr-1. The success of M. salmoides in Lake Chicamba was attributed to limited inter-specific and intra-specific competition for prey as a consequence of the timing of the spawning, cannibalism and high mortality rates.

Invasion of a headwater stream by non-native fishes in the Swartkops River system, South Africa.

In South Africa, fishes in headwater streams are increasingly being threatened by non-native fish invasions. These invasions originate primarily from source populations in mainstream rivers and impoundments. In the Blindekloof stream, a Swartkops River headwater tributary, the non-native Micropterus salmoides was eradicated from the stream by conservation authorities in 1989. As a result of this eradication subsequent occurrences of non-native fishes are most likely a result of upstream invasions from the mainstream Swartkops River. The Blindekloof stream therefore provided a unique opportunity to investigate whether M. salmoides would reinvade after its removal in 1989 and if the more recently introduced Clarias gariepinus and Tilapia sparrmanii would invade the Blindekloof stream. During snorkel and electrofishing surveys four non-native species were recorded (M. salmoides, M. dolomieu, T. sparrmanii, C. gariepinus). The large predators, M. salmoides, M. dolomieu and C. gariepinus were neither abundant nor widespread and T. sparrmanii was the only non-native species that had successfully established in the Blindekloof stream. Where M. salmoides and M. dolomieu were observed, all other fish species were absent. By contrast, the native species were present at all sites where C. gariepinus and T. sparrmanii were found. The Pseudobarbus afer population was predominantly limited to a 1.7 km section of the stream that was not invaded and is located above a waterfall. The limited distribution range of this Endangered species makes it particularly vulnerable to predation by non-native fishes. The results of this study indicate that these non-native fishes rapidly invaded the Blindekloof stream following their introduction into the mainstream.

Lake Malawi: fishes, fisheries, biodiversity, health and habitat

Lake Malawis fishes are a source of food for millions and provide a livelihood for thousands by encouraging tourism, fascinating the scientific fraternity, enchanting aquarists around the world and maintaining ecosystem processes in the lake. From a fisheries and resource assessment perspective, the region is data-poor, but there is sufficient peer-reviewed and grey literature on the limnology, fisheries and ichthyofauna of the lake to provide a good overview of the state of the fishery. There are signs of over exploitation and an increasing fishing effort has resulted in decreased catch rates, depletion of larger, more valuable species in the fishery and species changes. The fishery is harvesting stocks that were formerly thought to be under exploited. Previous attempts to manage the fishery have been ineffective and long term strategies addressing overfishing will need to transform the fishery from an open-access to a limited access system. As important as direct intervention in the management of the fisheries, will be the management of catchment processes. Increased nutrient inputs; changes to the phytoplankton composition; sediment loading; nearshore water quality impacts and changing water levels threaten the ecosystem. Introduction of alien invasive organisms is an ever present threat to the ecosystem as well, due to continued development of small scale aquaculture in the region. The overriding causative factor for all these effects is the poverty of the lakeshore communities which do not have the economic privilege of being able to adapt their utilisation patterns.

The biology of Tilapia rendalli and Oreochromis mossambicus (Pisces: Cichlidae) in a subtropical lake in Mozambique

This study of age and growth, reproduction and juvenile recruitment of Tilapia rendalli and Oreochromis mossambicus was undertaken in Lake Chicamba, a large, clear subtropical lake in central Mozambique. Sectioned otoliths were used to estimate age. Marginal zone analysis showed that annulus formation occurred during winter, in July and August. Maximum age estimates for T. rendalli and O. mossambicus were 16 and 10 years, respectively. Growth was described by the 3 parameter von Bertalanffy model as Lt = 238.74 (1-e-0,636(1 + 00905)) mm TL for T. rendalli and Lt = 266.06 (1-e-0 70904(1 + 0269)) mm TL for O. mossambicus. The length-at-50% maturity (Lm50) of male T. rendalli was 218 mm total lenglh (TL) and 205 mm TL in females. In O. mossambicus the L50 for males was 251 mm TL and for females 223 mm TL. Both species spawned throughout summer. Lake level was found to have no effect on spawning periodicity although recruitment was flood dependent.

Managing conflicts arising from fisheries enhancements based on non-native fishes in southern Africa.

Southern Africa has a long history of non-native fish introductions for the enhancement of recreational and commercial fisheries, due to a perceived lack of suitable native species. This has resulted in some important inland fisheries being based on non-native fishes. Regionally, these introductions are predominantly not benign, and non-native fishes are considered one of the main threats to aquatic biodiversity because they affect native biota through predation, competition, habitat alteration, disease transfer and hybridization. To achieve national policy objectives of economic development, food security and poverty eradication, countries are increasingly looking towards inland fisheries as vehicles for development. As a result, conflicts have developed between economic and conservation objectives. In South Africa, as is the case for other invasive biota, the control and management of non-native fishes is included in the National Environmental Management: Biodiversity Act. Implementation measures include import and movement controls and, more recently, non-native fish eradication in conservation priority areas. Management actions are, however, complicated because many non-native fishes are important components in recreational and subsistence fisheries that contribute towards regional economies and food security. In other southern African countries, little attention has focussed on issues and management of non-native fishes, and this is cause for concern. This paper provides an overview of introductions, impacts and fisheries in southern Africa with emphasis on existing and evolving legislation, conflicts, implementation strategies and the sometimes innovative approaches that have been used to prioritize conservation areas and manage non-native fishes.