Comparisons of isotopic niche widths of some invasive and indigenous fauna in a South African river

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Benthic assemblages of wetlands invaded by Tarebia granifera (Lamarck, 1822) (Caenogastropoda: Thiaridae) in the iSimangaliso Wetland Park,South Africa

Tarebia granifera is a freshwater/estuarine gastropod invading many tropical and sub-tropical areas around the world. This snail is native to southeast Asia and was accidentally introduced into South Africa during the last decade. The current study investigated shallow-water benthic assemblages of different invaded and uninvaded localities across locations spanning a large range of environmental conditions in the iSimangaliso Wetland Park. Using a correlation-based approach, we found that native benthic assemblages were more closely associated with environmental conditions than with densities of T. granifera. However, there were significant negative correlations between T. granifera abundance and Shannon Diversity at two of the invaded locations. This alien species has successfully invaded, and become dominant in, different types of water bodies with different assemblage compositions and physico-chemical characteristics, ranging from freshwater ponds to saline estuaries and lakes. The current data set is presented as an essential baseline for future studies. It is recommended that future work focuses on specific localities, in order to determine if changes in diversity are driven by non-native species or by other disturbances (e.g., climate change).     

Northern pike (<Emphasis Type="Italic">Esox lucius</Emphasis>) in a warming lake: changes in population size and individual condition in relation to prey abundance

The spring-fed Comal River in Texas, USA, has been impounded and channelized resulting mainly in a lentic environment with four headwater spring runs. We sampled two spring runs (lotic sites) and two lentic sites seasonally from April 2001 through April 2002 to assess (1) co-occurrence of native and exotic snail species (as determined by interspecific association), (2) the importance of habitat conditions in structuring relationships among these species, and (3) the distribution of snails infected with exotic trematode parasites. Three exotic and four endemic species of aquatic snails were collected, but only Elimia comalensis (Prosobranchia: Pleuroceridae, native), Melanoides tuberculatus, and Tarebia granifera (Prosobranchia: Thiaridae, exotic) were in sufficient densities for further analyses. Tarebia granifera was positively associated with both M. tuberculatus (χ² = 18.5, P < 0.001) and E. comalensis (χ² = 7.3, P < 0.01), although the co-occurrence between the two exotics was much stronger. Melanoides tuberculatus and E. comalensis exhibited a strong, negative association (χ² = 10.9, P < 0.001). The weaker co-occurrence between E. comalensis with the thiarids appeared to be driven by differences in habitat use by the thiarids and native E. comalensis. In lentic habitats, densities of M. tuberculatus and T. granifera were similar but differed significantly from E. comalensis whose densities were 200 times less than the exotic snails. In lotic spring runs, densities of T. granifera and E. comalensis were similar, but differed significantly from M. tuberculatus whose densities were 10 times fewer. Lower densities of M. tuberculatus and T. granifera in habitat conditions common to the spring runs may explain why exotic snail interactions were less with the native E. comalensis than with each other. The native snail, E. comalensis, was not infected with any trematodes, while 6.1% of M. tuberculatus and 4.8% of T. granifera were infected with exotic trematodes. Distributions of infected snails were aggregated; such that most infected snails were found in lentic habitats with silt substrates and moderate to high levels of detritus. Continued declines in spring-flows due to aquifer withdrawals and droughts will increase lentic habitats that may lead to increased densities of T. granifera and M. tuberculatus and their concomitant parasites. Handling editor: K. Martens     

Stable isotope evidence for dietary overlap between alien and native gastropods in coastal lakes of northern KwaZulu-Natal, South Africa

Background

Tarebia granifera (Lamarck, 1822) is originally from South-East Asia, but has been introduced and become invasive in many tropical and subtropical parts of the world. In South Africa, T. granifera is rapidly invading an increasing number of coastal lakes and estuaries, often reaching very high population densities and dominating shallow water benthic invertebrate assemblages. An assessment of the feeding dynamics of T. granifera has raised questions about potential ecological impacts, specifically in terms of its dietary overlap with native gastropods.

Methodology/Principal Findings

A stable isotope mixing model was used together with gut content analysis to estimate the diet of T. granifera and native gastropod populations in three different coastal lakes. Population density, available biomass of food and salinity were measured along transects placed over T. granifera patches. An index of isotopic (stable isotopes) dietary overlap (IDO, %) aided in interpreting interactions between gastropods. The diet of T. granifera was variable, including contributions from microphytobenthos, filamentous algae (Cladophora sp.), detritus and sedimentary organic matter. IDO was significant (>60%) between T. granifera and each of the following gastropods: Haminoea natalensis (Krauss, 1848), Bulinus natalensis (Küster, 1841) and Melanoides tuberculata (Müller, 1774). However, food did not appear to be limiting. Salinity influenced gastropod spatial overlap. Tarebia granifera may only displace native gastropods, such as Assiminea cf. ovata (Krauss, 1848), under salinity conditions below 20. Ecosystem-level impacts are also discussed.

Conclusion/Significance

The generalist diet of T. granifera may certainly contribute to its successful establishment. However, although competition for resources may take place under certain salinity conditions and if food is limiting, there appear to be other mechanisms at work, through which T. granifera displaces native gastropods. Complementary stable isotope and gut content analysis can provide helpful ecological insights, contributing to monitoring efforts and guiding further invasive species research.     

Introduction,distribution, spread,and impacts of exotic freshwater gastropods in Texas

We examined the patterns of distribution, vectors of introduction, and potential ecological impacts of freshwater exotic species in Texas over the last 45 years. Currently, five species of exotic gastropods are established: channeled-type applesnail (Pomacea insularum), red-rim melania (Melanoides tuberculatus), quilted melania (Tarebia granifera), giant rams-horn snail (Marisa cornuarietis), and Chinese mysterysnail (Cipangopaludina chinensis). In contrast to the northern part of the US, where shipping appears to be the most important vector for the introduction of aquatic invasive species, aquarium and ornamental trade dominated among unintentional vectors of introduction of all freshwater exotics in Texas, resulting in different patterns of distribution, spread, and ecological impacts. The rate of spread of exotic gastropods in Texas varied from 39 waterbodies colonized over 18 years for P. insularum to only three waterbodies during last 45 years for C. chinensis. Four of five exotic gastropods were found in highly vulnerable aquifer-fed springs and rivers, which contain numerous endemic and endangered species. The fifth species, Pomacea insularum, is an agricultural pest. Potential negative ecological effects of exotic gastropods include impacts on wetlands and wetland restoration, competitive exclusion of native snails, and the introduction of exotic parasites, trematodes, which could infect fish and waterfowl, including federally protected species. Aquifer springs with stable temperature regimes are refuges for both cold and warm intolerant species. Handling editor: D. Dudgeon     

Distribution patterns and habitat associations of Sandelia bainsii (Teleostei: Anabantidae), a highly threatened narrow-range endemic freshwater fish

Sandelia bainsii is a range-restricted and highly threatened freshwater fish endemic to South Africa. Recent genetic evidence suggests that this species comprises three allopatrically distributed lineages that have been informally designated as Sandelia sp. “Kowie,” Sandelia sp. “Keiskamma” and Sandelia sp. “Buffalo.” As these lineages have only been recently identified and are likely to face a high risk of extinction because of restricted distributions, there is a critical need for generating ecological information to guide conservation prioritisation. The present study compared the historical and current distribution patterns, together with the habitat associations of Sandelia sp. “Kowie” in the Koonap and Kat rivers, tributaries of the Great Fish River. This study indicated that this lineage has been extirpated from one of the three localities in the Koonap River where it was historically abundant. In the Kat River, the current distribution of Sandelia sp. “Kowie” was comparable to its historical range, but its future persistence is threatened by the presence of non-native piscivores, instream physical barriers and potential future exploration for shale gas and infrastructure development in the Karoo Basin. A generalised hurdle negative binomial model revealed that although this lineage's probability of occurrence was high in habitats with boulder and sand substrates, and low conductivity, habitat characteristics were poor predictors of its abundance. Thus, it was postulated that the current range of this lineage probably represents the only available habitats for the persistence of different life stages for this taxon. Alternatively, the observed patterns may suggest the possibility of a shift in habitat associations, possibly for optimum utilisation of the remaining refugia within this river system. Immediate conservation measures should focus on preventing the spread on non-native invasive fishes, whereas future studies should evaluate the impacts of population fragmentation and identify appropriate intervention measures to maintain this lineage's long-term adaptive potential.     

Water hyacinth,Eichhornia crassipes (Pontederiaceae), reduces benthic macroinvertebrate diversity in a protected subtropical lake in South Africa

The socio-economic impacts of the free-floating aquatic plant water hyacinth, Eichhornia crassipes (Pontederiaceae), on aquatic systems are well documented, yet the impacts on aquatic biodiversity, particularly invertebrate biodiversity, are less well understood. This study aimed to determine whether the presence of water hyacinth altered the diversity and assemblage structure of benthic macroinvertebrates in a conservation area. The benthic macroinvertebrate assemblage was sampled over 1 year at five sites under water hyacinth mats and at five sites without water hyacinth at Lake Nsezi—Nseleni River in the vicinity of Richards Bay, KwaZulu-Natal, South Africa. Artificial substrates were placed beneath water hyacinth mats or in the open water to allow for colonization by freshwater macroinvertebrates, and left for a period of 6 weeks, repeated on seven occasions. Twenty nine families comprising 18,797 individuals were collected, 817 (13 families) individuals were from under water hyacinth mat sites compared to 17,980 (27 families) individuals from open water sites. Ninety-eight percent of individuals collected were, however, the invasive snail, Tarebia granifera. Open water samples were separated from samples beneath the water hyacinth mat by non-metric Multidimensional Scaling, indicating reduced biodiversity associated with the presence of water hyacinth. Exclusion of the dominant Thiaridae from the analysis did not alter the groupings. Family richness(s) and abundance (N) were significantly higher in open water communities(S: H₃ = 21.09; P = 0.0001; N: H₃ = 22.58; P = 0.00001), while evenness (J’) was higher under water hyacinth mats (H₃ = 20.13; P = 0.0002). The presence of water hyacinth had a significantly negative impact on aquatic macroinvertebrate biodiversity in a conservation area, and therefore the control of this invasive aquatic plant must play a major role in catchment management.     

Black spot infection in juvenile steelhead trout increases with stream temperature in northern California

Climate change will increase water temperature in rivers and streams that provide critical habitat for imperiled species. Warmer water temperatures will influence the intensity and nature of biotic interactions, including parasitism. To better understand the factors influencing a neascus-type parasitic infection known as black spot disease, we examined the relationship between infection rate in juvenile steelhead trout (Oncorhynchus mykiss), abundance of another intermediate host (ramshorn snail, Planorbella trivolvis), and water temperature. We quantified infection patterns of trout at seven sites within the South Fork Eel River in northern California, visiting each site on three different occasions across the summer, and recording water temperature at each site. We also quantified infection patterns in trout captured from two tributaries to the South Fork Eel River. Overall, trout infection rates were highest in sites with the warmest temperatures. The abundance of ramshorn snails was positively related to both water temperature and black spot infection rates in juvenile trout. Both snail abundance and infection rates increased rapidly above a 23 °C daily maximum, suggesting a threshold effect at this temperature. We suggest that warmer temperatures are associated with environmental and biotic conditions that increase black spot disease prevalence in threatened steelhead trout. A comparison of our results with similar data collected from a more northern latitude suggests that salmonids in California may be warm-adapted in terms of their parasite susceptibility.     

Population structure of an invasive parthenogenetic gastropod in coastal lakes and estuaries of northern KwaZulu-Natal, South Africa

Background

Estuaries and coastal lakes receive little attention despite being heavily invaded by non-indigenous invasive species (NIS). In these situations, studies of population dynamics in invaded habitats can provide valuable insights into how NIS interact with new environments. Tarebia granifera is a prosobranch gastropod from south-east Asia which has invaded other sub-tropical parts of the world. This study addresses whether a small number of key environmental factors influences gastropod communities, and specifically how the population density and size structure of T. granifera were influenced by environmental change in estuaries and coastal lakes in southern Africa.

Methodology/Principal Findings

T. granifera''s density, number of brooded juveniles and size structure were measured at the St. Lucia Estuary, Mgobozeleni Estuary, Lake Sibaya and Lake Nhlange. Size structure was classified according to shell height (SH). All dissected individuals were found to be female and free from trematode infection. Salinity, water depth, temperature, and pH were the main factors correlated with population density of gastropod communities. T. granifera often reached densities well over 1000 ind. m⁻², displacing indigenous gastropods and becoming a dominant component of the benthic community. T. granifera successfully invaded estuaries despite frequent exposure to high salinity and desiccation, which could together eliminate >97% of the population. The persistence of T. granifera was ensured due to its high fecundity and the environmental tolerance of large adults (20–30 mm SH) which carried an average of 158±12.8 SD brooded juveniles. Repeat introductions were not essential for the success of this parthenogenetic NIS.

Conclusion/Significance

There is a need for a broader study on the reproductive biology of T. granifera (including the previously overlooked “brood pouch ecology”), which affects population dynamics and may be relevant to other parthenogenetic NIS, such as Melanoides tuberculata and Potamopyrgus antipodarum.     

Record of Blue tilapia Oreochromis aureus (Steindachner, 1864) in the Eerste River catchment,Western Cape province,South Africa

Oreochromis aureus was imported from Israel into South Africa in 1959 but data on its current status in South Africa are lacking. Genomic DNA was extracted and the COI gene amplified at the South African Institute for Aquatic Biodiversity. The identity of the sequences and specimens was determined using the Barcode of Life Data Systems and GenBank. Morphological and genetic assessment demonstrated that 11 specimens collected from two farm dams in the Eerste River System, Western Cape province, were Oreochromis aureus. A MaxEnt model compiled using global distribution, rainfall and temperature data predicted that large areas of southern Africa were climatically suitable for this species, indicating considerable invasion debt in southern Africa. As a result, surveys to assess for the extent of the invasion in South Africa and eradication of existing populations, if feasible, are recommended management actions.     

Detrital Fruit Processing in a Hawaiian Stream Ecosystem1

In temperate forested streams, fruit from riparian trees is generally a minor and seasonal component of the allo‐chthonous detritus. In contrast, riparian fruit input to tropical streams is often high and continuous. Detrital fruit is abundant in some forested Hawaiian streams compared to other forms of riparian detritus, and rates of leaf litter processing by macroscopic invertebrates are very low. These observations suggested that fruit is an important food resource for detritivores. A microcosm system was used to measure the rates at which two common detrital fruits, guava and mango, were processed by two common detritivores, the prawn Macrobrachium lar and the gastropod Tarebia granifera. Comparisons of fruit weight loss rates normalized by detritivore weight indicated that M, lar processed guavas at significantly higher rates than T. granifera, differences in rates of mango processing by M. lar and T. granifera were not significant. Microcosms containing both M. lar and T. granifera were used to test for interactions between the invertebrates that affected rates of mango processing. No interspecific interactions were detected. A field study was conducted in Kaiwiki Stream, Island of Hawaii, to determine rates of detrital fruit input and export. Detrital fruit was supplied to the study area year‐round, with peaks corresponding to summer and autumn fruiting seasons. Guavas and mangos accounted for 85 percent of the fruit biomass entering the stream and 92 percent of the fruit exported from the stream. Mean daily export rates of guava were 7 percent of input, and export rates of mango were 5 percent of input. These measurements suggested that most of the fruit entering the stream is retained and comprises a substantial food resource for detritivores. Comparisons of the biomass‐specific rates at which M. lar and T. granifera processed mangos and guavas with the rates at which mangos and guavas entered Kaiwiki Stream suggested that these invertebrates can process most of the detrital fruit in the stream.     

The phylogeny of the genus Indoplanorbis (Gastropoda,Planorbidae) from Africa and the French West Indies

Indoplanorbis exustus is a freshwater snail known as the intermediate host of various trematode parasites, including different species of the genus Schistosoma. Although its genetic diversity is well described in Asia, the phylogenetic diversity of strains from Africa and Guadeloupe (French West Indies) and their relationship to Asian and South‐East Asian strains remain unknown. To tackle this issue, we sampled individuals from Africa and Guadeloupe, and we computed phylogenetic reconstructions using five molecular markers: partial sequences of two mitochondrial genes, cox1 and 16S, and three nuclear markers, ITS1, ITS2 (Internal Transcribed Spacer 1 and 2) and 5.8S. Our results suggest that strains in Africa and Guadeloupe come from Asia and that they all belong to a single clade that is widespread around the globe.     

Light,temperature and flow regimes of the Vaal River at Balkfontein,South Africa

A detailed investigation into light, temperature and flow regimes of the Vaal River was done for the first time. The Vaal River is ecologically and economically one of the most important rivers in South Africa. In a South African context, the Vaal River, with an average Secchi disk depth (Z _sd) of 0.41 m and an average euphotic zone depth (Z _eu) of 1.3 m, is a moderately turbid system. Red light (approx. 670 nm) penetrated Vaal River water to a greater degree than other components of the light spectrum. Water temperature in the Vaal River (min. = 10, max. = 27 °C at midday), closely followed seasonal changes in average atmospheric temperatures. The water-column shows little temperature stratification. The Vaal River is a highly regulated system with relatively high discharge rates at Balkfontein (average 112 m³ s^–1), but relatively low current velocity (approx. 0.35 m s^–1). Discharge was the most important variable to influence transparency of Vaal River water. Higher discharge resulted in higher total suspended solids (TSS) concentration and higher extinction coefficient (k) values as well as in higher turbidity and thus in lower Z _eu and Z _sd. From the mean TSS (141 mg l^–1) it was calculated that the Vaal River (mean annual run-off of 3532 million m³) transported 498016 tonnes of suspended solids per annum.     

NEW DISTRIBUTION RECORD FOR CHILOGLANIS BIFURCUS JUBB AND LE ROUX, 1969 IN A TRIBUTARY OF THE CROCODILE RIVER,MPUMALANGA PROVINCE,SOUTH AFRICA

Summary

A new distribution record for the South African Red Data fish species Chiloglanis bifurcus was found in the Nels River at an altitude of 830 m a.m.s.l. in the Mpumalanga province, South Africa. Previously this species was known to occur between altitudes 900 m and 1200 m a.m.s.l. in the Elands and Crocodile Rivers of the Inkomati River system, as well as in tributaries of these rivers, namely the Gladdespruit, Stadspruit and Ngodwana River. It was also found to occur in the Komati River of the same river system.     

1. BIRD-LIFE AT THE ORANGE RIVER MOUTH

Kalejta, B. 1993. Diets of shorebirds at the Berg River estuary, South Africa: spatial and temporal variation. Ostrich 64: 123–133.

The diets of three common migrant waders; Curlew Sandpiper Calidris ferruginea. Grey Plover Pluvialis-squatarola and Greenshank Tringa nebularia and three resident species, Blacksmith Vanellus armatus, Kittlitz's Charadrius pecuarius and Whitefronted Plovers C. marginatus, were studied at the Berg River estuary, South Africa from December 1987 to April 1989. Direct observations of feeding were combined with analyses of stomach contents, pellets and droppings. Nereid worms, Ceratonereis erythraeensis and C. keiskama, were the principal food of all species studied except Greenshanks, which fed mostly on crabs, Hymenosoma orbiculare. Despite considerable overlap in the diets of all bird species, there were differences in the size classes of nereids taken by different bird species. Visually-foraging plovers (Charadriidae) were highly selective, feeding on the largest nereids regardless of their abundance. Tactile foragers, represented by Curlew Sandpipers, were non-selective, and consumed different size-classes of worms in proportion to their abundance in the substratum. Seasonal variations in the diet of waders are linked to seasonal changes in the availability of prey species.     

DNA bar-coding reveals source and patterns of <Emphasis Type="Italic">Thaumastocoris peregrinus</Emphasis> invasions in South Africa and South America

Thaumastocoris peregrinus is a recently introduced invertebrate pest of non-native Eucalyptus plantations in the Southern Hemisphere. It was first reported from South Africa in 2003 and in Argentina in 2005. Since then, populations have grown explosively and it has attained an almost ubiquitous distribution over several regions in South Africa on 26 Eucalyptus species. Here we address three key questions regarding this invasion, namely whether only one species has been introduced, whether there were single or multiple introductions into South Africa and South America and what the source of the introduction might have been. To answer these questions, bar-coding using mitochondrial DNA (COI) sequence diversity was used to characterise the populations of this insect from Australia, Argentina, Brazil, South Africa and Uruguay. Analyses revealed three cryptic species in Australia, of which only T. peregrinus is represented in South Africa and South America. Thaumastocoris peregrinus populations contained eight haplotypes, with a pairwise nucleotide distance of 0.2–0.9% from seventeen locations in Australia. Three of these haplotypes are shared with populations in South America and South Africa, but the latter regions do not share haplotypes. These data, together with the current distribution of the haplotypes and the known direction of original spread in these regions, suggest that at least three distinct introductions of the insect occurred in South Africa and South America before 2005. The two most common haplotypes in Sydney, one of which was also found in Brisbane, are shared with the non-native regions. Sydney populations of T. peregrinus, which have regularly reached outbreak levels in recent years, might thus have served as source of these three distinct introductions into other regions of the Southern Hemisphere.     

Changes in the algal composition and water quality of the Sundays River,Karoo, South Africa,from source to estuary

A first survey was done on algae present in the Sundays River from its source to its confluence with the sea. Species found in the upstream sections of the river included indicators of good water quality, but the quality deteriorated downstream with peaks in algal abundance being ascribed to peaks in nutrient concentrations. Cyanobacteria and euglenoids were present in the upper and middle reaches of the river, but were absent downstream. Dinoflagellates became more important downstream, especially in the estuary. Dominant species, reaching high concentrations along the river, included Nitzschia frustulum, Nitzschia capitellata, Carteria klebsii, Chlorella vulgaris and Anabaena species. The presence of the diatom Eolimna comperei is a first record of its occurrence in South Africa. The Sundays River can be described as a brackish, hard water system with high nutrient concentrations in certain sections. The most important contributors to high nutrient concentrations were point sources in the vicinity of towns along the river banks, as well as diffuse sources contributing to high nitrogen concentrations in the fertile Sundays River valley. Increasing salinities were due to pollution, evaporation and agricultural activities in the valley.     

Record of Eldana saccharina Walker (Lep,Pyralidae) in inland South Africa and its genetic relationship with the coastal population

Surveys to investigate the distribution and abundance of stem borers in natural habitats were conducted in February 2006 and January–February 2007. The surveys included eastern, northern and central parts of South Africa as well as three localities in Lesotho. During the surveys, Eldana saccharina Walker was recovered from three new localities in inland South Africa and two new indigenous hosts, Phragmites australis Cav. and Panicum maximum Jacq. from Boskop in the North‐West Province and Oribi Gorge in KwaZulu‐Natal respectively. Populations of E. saccharina in different parts of Africa are known for their differences in larval feeding behaviours, host plant choice and natural enemies. It is important to understand the origin of the newly recovered population for prevention of incursion and efficient management in case it invades crops. Molecular analysis indicated that the populations recovered in these new locations and from the new host plants are part of the southern African population of E. saccharina. With change in climate, and disturbance in wetlands the insect is expected in the future to be more abundant and problematic in inland areas of southern Africa.     

Portulacaria afra is constrained under extreme soil conditions in the Fish River Reserve, Eastern Cape, South Africa

Restoration of degraded thicket landscapes can be achieved by planting Portulacaria afra (spekboom) cuttings. The factors determining P. afra abundance in thicket types not dominated by P. afra are of interest to restoration practitioners because they could influence restoration protocols using P. afra cuttings in these landscapes. We consequently investigated the relationship between P. afra cover and soil properties, namely pH, EC, organic C, particle size distribution and total content of 26 elements at 78 sites in the Fish River Reserve, Eastern Cape province, South Africa. P. afra cover showed a consistent pattern of constraint at extreme levels (both high and low) of subsoil EC, organic C, sand content, Ca, Zn and Al. The results suggest that P. afra is most competitive in intermediate edaphic environments in the Fish River Reserve. It remains to be investigated in restored landscapes whether P. afra cutting survivorship and growth are greater in such soils.     

Using a unified invasion framework to characterize Africa’s first loricariid catfish invasion

This paper presents evidence of establishment of a loricariid population in the Nseleni River in South Africa and uses a unified framework to determine its invasion stage. Specimens were identified morphologically as Pterygioplichthys disjunctivus (Weber 1991), but genetic barcoding results indicated close association with specimens that may have a hybrid history. The species was introduced into South Africa via the pet trade and the first record of introduction into the wild was in 2004. Samples collected in 2011 and 2012 demonstrated that there were multiple length cohorts in the population including juveniles (12–130 mm total length TL) and large (>300 mm TL) adult fish. Gonadal assessment of adults demonstrated the presence of reproduction capable specimens. The concurrent occurrence of mature adults and juvenile fish demonstrated establishment. Locality records indicate that P. disjunctivus has already spread between two rivers through an inter basin water transfer. Using a unified framework for invasions this invasion was categorized as a self-sustaining population in the wild with individuals surviving and reproducing a significant distance from their original point of introduction. Containment is suggested as potential management strategy.