Bruce R. Ellender

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.

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.

Validating and corroborating the deposition of two annual growth zones in asteriscus otoliths of common carp Cyprinus carpio from South Africa's largest impoundment

A total of 816 common carp Cyprinus carpio asteriscus otolith pairs were collected from Lake Gariep, South Africa. Otoliths were interpreted whole, submerged in methyl salicylate and viewed under transmitted light. The precision of growth zone counts of the primary reader was estimated at 5·54 and 7·03% using the average per cent error method and the coefficient of variation, respectively. Age-bias plots indicated no systematic bias between the primary reader and the three secondary readers for up to nine growth zones (95% of the sample). Growth zone deposition rate was validated using a mark-recapture experiment of chemically tagged C. carpio (n = 21) conducted in a large earthen pond under ambient conditions in the vicinity of Lake Gariep. The validation results were corroborated for the wild population by edge analysis and a length-based age-structured model. All three methods suggest that growth zone formation occurred biannually, exemplifying the importance of age validation as a prerequisite for understanding the life history of C. carpio.

Fish distributions in the Rondegat River, Cape Floristic Region, South Africa, and the immediate impact of rotenone treatment in an invaded reach

Alien fishes are considered the most serious threat to native headwater stream fishes in South Africa. A 4 km reach of the Rondegat River is the first section of a South African river to be rehabilitated through the attempted removal of alien fish by using the piscicide rotenone. The objectives of the current study were to establish the distribution and relative abundance of native and alien fish prior to treatment, and to assess the immediate impact of the treatment on the fish population. Forty-three sites were sampled using backpack electrofishing, snorkel transects and underwater video analysis. In the invaded lower reaches, native Labeobarbus capensis was detected only at very low densities, while three other native fish species were not detected. Alien fish were not detected above a barrier waterfall 5 km upstream of the rivers confluence with a reservoir. The fish density of 97 fish per 100 m2 in non-invaded reaches was more than an order of magnitude higher than that of 7 fish per 100 m2 in the invaded reach. A total of 470 Micropterus dolomieu and 139 L. capensis were removed from a 4 km treatment zone during the rotenone operation. No fish were detected in this area after the rotenone treatment.

Validation of Growth Zone Deposition in Otoliths of Two Large Endemic Cyprinids in Lake Gariep, South Africa

We tested the hypothesis that growth zones in the astericus otoliths of smallmouth yellowfish (Labeobarbus aeneus) and Orange River mudfish (Labeo capensis) were deposited annually, Two methods, fluorochrome marking and edge analysis of otoliths were used. For fluorochrome marking, specimens of both species were injected with 60 mg/kg fish mass Oxytetracycline hydrochloride (OTC) and released into large earthen ponds under ambient conditions adjacent to Lake Gariep. Twenty-three L. aeneus and one L. capensis were recaptured 10–14 months later. Edge analysis was based on the optical interpretation of L. aeneus (n = 342) and L. capensis (n = 512) otolith margins collected between November 2006 and May 2008 from Lake Gariep. The frequency distribution of opaque margins over time was fitted using a binomial periodic regression. The estimated cycle length was not significantly different from a hypothesized 12 months for both species. The number of growth zones distal to the OTC mark was consistent with findings from the edge analysis, providing evidence that growth zones in astericus otoliths of both species can be interpreted as annuli.

Using functional responses to quantify interaction effects among predators

Summary Predator diversity alterations have been observed in most ecosystems as a result of the loss and/ or addition of species. This has implications for predator–prey dynamics as non-trophic interactions among predators, so-called multiple predator effects (MPE), are known to influence predation success. In addition, there is often a density-dependent relationship between prey availability and prey consumption (functional response). While MPE investigations are common in the literature, functional responses have rarely been incorporated into this field of predation ecology. Here, we outline an experimental procedure that incorporates functional responses into multiple predator effect studies. Using three fish species with different functional traits as model predators (bluegill Lepomis macrochirus, southern mouthbrooder Pseudocrenilabrus philander and banded tilapia Tilapia sparrmanii), we assess intra- and interspecific predator interaction outcomes on predator–prey dynamics. This was done by contrasting observed functional responses of heterospecific and conspecific combinations of predators with expected responses based on those of individual predators. Multipredator combinations produced variable results. Bluegill were the only species in which observed conspecific multipredator functional responses matched those of expected based on individual performance (prey risk neutral effects). In contrast, prey risk reduction was observed for both mouthbrooder and tilapia conspecific multipredator trials. Heterospecific combinations revealed strong prey risk reduction effects for mouthbrooder–tilapia and bluegill–tilapia trials, while mouthbrooder–bluegill multipredator functional responses combined additively. These results are discussed within the context of behavioural traits of the species and the development of a trait-based predictive framework. Using a functional response approach allowed for the assessment of multiple predator effects across a range of prey densities. We propose that the incorporation of within-guild predator combinations into classic functional response investigations will enhance predictive capacity development in competition and predation ecology. A lay summary is available for this article.

Life history and population dynamics of invasive common carp, Cyprinus carpio, within a large turbid African impoundment

As one of the most widely established freshwater fishes globally, the invasive success of common carp, Cyprinus carpio, is beyond dispute. Although detailed knowledge on its biology would assist in its management, relatively few life history studies have investigated C. carpio outside of its natural range. The present study analyses the life history and population structure of exotic C. carpio in a 364 km2 African impoundment. We used whole astericus otoliths to show that C. carpio attained ages of up to 7 years and grew more rapidly than has been recorded for any population from Europe or Australia. Macroscopic staging of gonads indicated protracted spawning, with highest reproductive activity observed during late spring and early summer. Total mortality, natural mortality and fishing mortality rates were estimated at 0.72, 0.60 and 0.12 year–1 respectively. Life history comparisons across the species’ global distribution showed large variations in growth and longevity, whereas early maturation (2–3 years) appeared to be a rather conservative trait. The combination of early maturity, fast growth and relatively high natural mortality rates suggests a rapid population growth potential associated with high resilience to management interventions once C. carpio has been introduced.

Age and growth and maturity of southern Africa's largest cyprinid fish, the largemouth yellowfish Labeobarbus kimberleyensis.

The aim of this study was to use specimens of the largemouth yellowfish Labeobarbus kimberleyensis, southern Africas largest cyprinid [IUCN red-listed as Near Threatened (NT)], obtained from gillnet by-catch to describe aspects of its biology in order to assist future conservation and management decisions. Ninety three L. kimberleyensis were collected between March 2007 and May 2008 from Lake Gariep, South Africa. Labeobarbus kimberleyensis was present in 38% of all gillnet catches, but in low numbers (2% of the catch) and it contributed 8% to the catch by mass. Age was estimated using astericus otoliths. Growth increment formation on these otoliths was validated as annual using edge analysis and the mark-recapture of chemically tagged captive fish. Resultant analysis showed that the species is slow growing and the oldest aged fish was a 17 year, 690 mm fork length (L(F)) male. The smallest ripe female fish measured 394 mm L(F) and was 7+ years old and the smallest mature male was 337 mm L(F) and 5+ years old. Slow growth and late maturity make this species vulnerable to exploitation emphasizing the need for continued high conservation priority.

Juvenile population dynamics of Oreochromis mossambicus in an intermittently open estuary at the limit of its natural distribution

ABSTRACT Juvenile Oreochromis mossambicus in the 25 ha, temperate, intermittently open, East Kleinemonde Estuary in the Eastern Cape, South Africa (33°32′S, 27°03′E), showed a seasonal shift in distribution, occupying all areas of the estuary in summer and, subsequently, being limited to the upper reaches in winter. The young of the year in the littoral zone were rapidly depleted and resultant estimates of the instantaneous rate of total mortality (Z) ranged from 5.4/yr to 6.9/yr. High annual rates of juvenile mortality and small adult populations imply that stock recovery after a reduction in adult population numbers, through activities such as fishing, may be slow. This work adds to the knowledge base of O. mossambicus ecology and population dynamics in temperate estuaries at the limit of its natural distribution

Trophic overlap between fish and riparian spiders: potential impacts of an invasive fish on terrestrial consumers.

Abstract Studies on resource sharing and partitioning generally consider species that occur in the same habitat. However, subsidies between linked habitats, such as streams and riparian zones, create potential for competition between populations which never directly interact. Evidence suggests that the abundance of riparian consumers declines after fish invasion and a subsequent increase in resource sharing of emerging insects. However, diet overlap has not been investigated. Here, we examine the trophic niche of native fish, invasive fish, and native spiders in South Africa using stable isotope analysis. We compared spider abundance and diet at upstream fishless and downstream fish sites and quantified niche overlap with invasive and native fish. Spider abundance was consistently higher at upstream fishless sites compared with paired downstream fish sites, suggesting that the fish reduced aquatic resource availability to riparian consumers. Spiders incorporated more aquatic than terrestrial insects in their diet, with aquatic insects accounting for 45–90% of spider mass. In three of four invaded trout rivers, we found that the average proportion of aquatic resources in web‐building spider diet was higher at fishless sites compared to fish sites. The probability of web‐building and ground spiders overlapping into the trophic niche of invasive brown and rainbow trout was as high as 26 and 51%, respectively. In contrast, the probability of spiders overlapping into the trophic niche of native fish was always less than 5%. Our results suggest that spiders share resources with invasive fish. In contrast, spiders had a low probability of trophic overlap with native fish indicating that the traits of invaders may be important in determining their influence on ecosystem subsidies. We have added to the growing body of evidence that invaders can have cross‐ecosystem impacts and demonstrated that this can be due to niche overlap.