Review Article |
Academic editor: Shana McDermott
© 2024 Calum MacNeil, Robin Holmes, Edward Challies, Kiely McFarlane, Jason Arnold.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
MacNeil C, Holmes R, Challies E, McFarlane K, Arnold J (2024) Strangers in a strange land; freshwater fish introductions, impacts, management and socio-ecological feedbacks in a small island nation – the case of Aotearoa New Zealand. NeoBiota 94: 101-125. https://doi.org/10.3897/neobiota.94.122939
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Invasive non-native species (INNS) are key drivers of global biodiversity loss. This is particularly evident in freshwater ecosystems, where the rates of both vertebrate biodiversity loss and biological invasion exceed those of marine and terrestrial systems. Aotearoa New Zealand (henceforth Aotearoa) like many other island nations, has a troubled history with NNS. However, it is also unique, as the main islands were the last major landmasses on Earth to remain uninhabited by humans. The endemic fauna had evolved in isolation from any anthropogenic influence or introduced NNS, until the mid-thirteenth century with the arrival of Māori, the first people to inhabit Aotearoa. Centuries later, following European colonisation, many non-native freshwater fish were deliberately introduced by acclimatisation societies. Currently, most of the native freshwater fish species of Aotearoa are at risk of extinction, despite almost 90% of these being found nowhere else on earth. Many of these species are highly valued by the indigenous people of Aotearoa, who have repeatedly highlighted biases towards NNS in freshwater fish management. With the rate of biological invasions increasing, it is timely to address interconnected issues concerning the history, impacts, management and current / future policy directions, including those involving biosecurity, for non-native freshwater fish in Aotearoa. We do this by applying a social-ecological systems (SES) lens, with a focus on causal-loop relationships and feedbacks to improve understanding of the dynamics of drivers, mechanisms and impacts of such invasions. We highlight the tensions that have resulted from managing some NNS as ‘pests’ threatening native biodiversity, while simultaneously promoting a tourism and recreational fishery resource for specific NNS. This has generated extremely polarized views on the ‘status’ of non-native freshwater fish species and given rise to contradictory and divergent goals for their management. We show how a disjointed and often incoherent policy landscape has contributed to legal ‘anomalies’ for NNS, including policy misalignments and gaps, hampering effective use of resources, while also entrenching contradictory management programmes for different stakeholders. Our study shows how these interconnected issues have been manifested in social-ecological feedback loops on core aspects of NNS policy and management, past and present. Consequently, there is a need for increased comprehension of the diverse array of potential impacts of NNS for different environments, stakeholders and Māori while developing coherent and practical management methods to reduce such impacts and improve social-ecological resilience. We conclude that adopting a SES approach will aid this endeavour.
Biosecurity, causal-loops, fish introductions, freshwater fish policy, non-native species, social-ecological system
Invasive non-native species (INNS) are intrinsically linked with many stakeholder actions (governments and public) throughout the world, to affect human wellbeing via negative or positive social-economic and social-ecological impacts. An INNS is a non-native species (NNS) which has expanded its range beyond its point of introduction and has had a measurable impact on factors such as ecology, ecosystem services, economics and human health (
Aotearoa New Zealand (henceforth Aotearoa) like many other island nations, has had a troubled history with NNS, stretching back several centuries (
The historical context of biological invasions and introductions in Aotearoa is globally unique (
Although there is a complex interplay among drivers of freshwater fish invasions (
Different stages of biological invasion (adapted from
Current Aotearoa policy for in-stream structures such as fish passages to simultaneously act as barriers to ‘undesirable’ species spread and facilitators of ‘desirable’ species spread focussed through human-natural systems and simultaneous negative (mitigating) and positive (reinforcing) feedback loops for ‘undesirable’ and ‘desirable’ species respectively.
These feedbacks provide a useful starting point from which to build a SES approach to understand interdependent systems, such as governance institutions and goals for development, biodiversity and ecosystem services (
The identification and strengthening of negative feedback loops associated with biosecurity, is also increasingly important in an era where climate change is accelerating the rate of biological invasion (
Over recent decades, concerns about the ecological impacts of freshwater NNS have existed alongside a drive for economic expansion of aquaculture and sports-fishing / angling tourism, shaping government policies. The tensions that have resulted from managing some NNS as ‘pests’ threatening native biodiversity, while simultaneously promoting a tourism / fishery resource for other NNS, have unsurprisingly generated highly polarized views on the ‘status’ of introduced freshwater fish species (
It is increasingly argued that there is a need to ‘strike the right balance’ between native and non-native species in fisheries management in Aotearoa (
A social-ecological systems (SES) approach can provide insights into the drivers, mechanisms and impacts of biological invasions (
To understand why freshwater fish legislation has evolved as it has, it is first important to appreciate the impacts of past and current invasions documented in Aotearoa, and second, to consider the risk profile of future potential invaders, which have had impacts in similar bioclimatic regions, albeit with different faunal assemblages (
Such fish introductions can drive positive socioeconomic feedback loops that increase donor fish transfer out of source regions over time. This is because the perceived success and value of previous fish introductions in recipient areas increases demand, ‘trade volume’ and the number of potential species within these trades (
Socio-ecological invasion meltdown (positive, reinforcing feedback loop) for brown trout introductions into the British colony of New Zealand by Victorian acclimatisation societies. Intentional introduction and establishment in recipient regions is done without consideration of any potential negative impacts on native fish species and the cultural / resource values that the indigenous human population attribute to these.
The active ‘acclimatisation’ movement continued up until as recently as the late 1960s in Aotearoa (
However, even in Victorian times there was some questioning of the general lack of appreciation of the impacts of introduced species on the receiving environment. The author of a letter to The Colonist newspaper in 1873 calling themselves a ’Disbeliever in Too Much Acclimatisation’, jokingly suggested that if leopards were introduced, they could be sustained on local school children (
Introduced freshwater fish can have a wide variety of direct and indirect ecological impacts on native fish and ecosystems (for examples of major reviews see
Although the brown trout is by far the best studied non-native fish in terms of ecological impacts, it is just one of many freshwater fish introductions that have impacted waterways in Aotearoa. Previous global reviews of the ecological impacts of freshwater non-native fish species have found it helpful to consider impacts at different levels or scales of biological organization, ranging from genetic to individual to population to community and finally ecosystem level (
Examples of ecological impacts of non-native freshwater fish species in Aotearoa. (Note only Aotearoa based studies are included and potential effects ascribed to overseas studies are purposely excluded).
Scale of biological organization | Impact and examples of mechanisms | Description |
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Genetic | Altering genetic resources via hybridization. | Tench may have the potential to hybridize with other introduced cyprinid fish such as goldfish, rudd and orfe ( |
Individual | Animal health and growth via altered behaviour, disease and parasite transmission. | Diel rhythm of habitat (water column and stream substratum) use by mayfly nymphs was affected by presence of brown trout, which exerted a different selection pressure on invertebrate drift behaviour than native galaxiids ( |
Tench have a potentially high infestation rate with the gut parasite Ligula intestinalis, which is already present in North Island lakes. Tench populations could act as a reservoir for this parasite which also infects native fish such as common bullies ( |
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Interactions between mosquitofish and native mudfish (Galaxiidae) can negatively impact mudfish foraging behaviour and prey capture rates ( |
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Population | Population size decline via predation, competition, disease and parasite transmission. | Brown trout have replaced nonmigratory galaxiid fish in some streams and diminished population sizes in others and have altered the distribution / range of large invertebrates such as crayfish ( |
Brown trout and rainbow trout feed on a range of macroinvertebrate species in a South Island lake ( |
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Predation by rainbow trout has caused significant declines in koaro (a native galaxiid fish) in North Island lakes ( |
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Small brown bullhead catfish feed on chironomids, Cladocera, gastropods, caddisfly larvae, plant material and detritus. and large catfish prey on native crayfish, fish and terrestrial invertebrates in Lake Taupo ( |
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Perch include smelt, common bullies and macroinvertebrates such as mysids and damselflies in their diets in a South Island river ( |
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Mosquitofish predate fry of native black mudfish and may exclude them from some habitats ( |
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In lake field trials and tank experiments rudd selectively eat different macrophytes, potentially influencing composition of macrophyte communities and this selective feeding may prevent re-establishment of these species in restoration programmes ( |
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An intensive removal of koi carp from a North Island lake led to significant reductions in the koi carp population, which coincided with an increase in native eel abundance, suggesting dietary overlap and competition for food between carp and eels may have been suppressing eel population size ( |
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Tench populations could act as a reservoir for a gut parasite and widespread transmission could reduce populations of native fish hosts such as common bullies ( |
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Community | Species extinction and reduction in native biodiversity | Presence of brown trout is a driver for elimination of non-migratory galaxiids from some streams ( |
Changes in composition of native species assemblages. | Brown trout have suppressed grazing pressure from macroinvertebrates on algae biomass and thus can enhance algal biomass and alter algal species composition and potentially macroinvertebrate community ( |
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Alteration / modification of food webs. | Annual production of macroinvertebrates is consumed by brown trout and not galaxiids where the native fish have been replaced and algal primary productivity can be six times higher in a ‘brown trout stream’ than a ‘galaxiid stream’ ( |
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Mosquito fish (Gambusia affinis) can induce changes in zooplankton community which diminish diets of native mudfish ( |
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Ecosystem | Modification of nutrient cycles. | Greater algal primary productivity in streams where brown trout have replaced galaxiids leads to an increased nutrient flux from the water to benthic community ( |
A range of NNS (rudd, tench, perch, brown bullhead catfish, goldfish, and koi carp) alters nutrient levels, affecting lake trophic processes, with excretion and bioturbation increasing nutrient levels ( |
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Loss / modification of habitat, native species refuges. | Salmon redd construction by Chinook salmon in two salmon spawning streams decreased the abundance of mosses, algae, macrophytes and sediment and detritus, causing a geomorphic modification of pool-riffle sequences and this was associated with a decrease in the abundance of benthic macroinvertebrates ( |
It must also be appreciated that any consideration of ecological impacts needs to acknowledge the complicated and multifaceted mechanisms at play across different spatial and temporal scales. Thus, the various impacts on native species and ecosystems described in Table
Nine non-native freshwater fish species have been identified as the most serious ‘pests’ for natural heritage managers in Aotearoa in terms of their ecological impact and spread (
Table
At its simplest level, biosecurity consists of a number of feedback loops (positive and negative) aiming to influence different aspects of the biological invasion process (Fig.
If we focus on fisheries management in countries such as Aotearoa, government resources and legislation are invariably applied to a limited suite of species deemed ‘important’, whether these are ‘desirable’ natives or non-natives which policies favour, or ‘undesirable’ non-natives which policies work against (see
Biosecurity, specifically as it relates to non-native freshwater fish in Aotearoa, is regulated in the context of four principal pieces of legislation: the
Prevention is always better than cure when it comes to biological invasion and the Hazardous Substances and New Organisms Act 1996 (
A core aspect of any biosecurity programme requiring public support is the language used to describe species and biosecurity objectives. ‘Branding’ certain species as ‘invasive aliens’ or as ‘pests’, sends out an unambiguous message, that such species are undesirable (
Pest, alien, invasive, unwanted, undesirable and noxious are all terms that government agencies have purposely used to prompt the public to actively discriminate against non-native species labelled as such (
Main government legislation in Aotearoa dealing with the biosecurity, presence / possession and culture of non-native freshwater fish species and the classifications of selected fish species that have resulted from application of such legislation (table adapted from
While this multiple labelling could be regarded as a pragmatic fisheries management approach, we suggest that it also reflects problems with policy (such as duplication), which are exacerbated by a current lack of ‘joined up’ centralised and holistic management of freshwater fisheries. Currently, different sectors of government, as well as public entities, have different responsibilities and accountabilities for different freshwater fisheries. To complicate matters further, a core piece of freshwater environmental legislation, the National Policy Statement for Freshwater Management (NPSFM-2020) only explicitly protects the habitats of two freshwater fish species and these are both NNS, namely trout and salmon. Perversely, only one native freshwater fish has full legal protection in Aotearoa legislation (the
Members of the Carp family (Cyprinidae) are also subject to multiple labels in Aotearoa, depending on the species involved and their perceived ‘usefulness’ or threat. The koi carp is classified as both a noxious fish and an unwanted organism, depending on the legislation applied (
While legal approval must be sought to possess a range of named freshwater animal and plant pests, including non-native Gambusia mosquito fish, no approvals at all are needed for a species such as the goldfish (Carassius auratus). Indeed, there are currently many freshwater NNS that are not covered by existing legislation and therefore effectively have no legal status in the context of current legislation. Apart from goldfish, these NNS ‘gaps’ with no legal status, include other aquarium / ornamental fish, brown bullhead catfish (Ameiurus nebulosus) and invertebrates. Although catfish and goldfish have no strict legal status, fishing regulations for recreational and commercial fishers require all captured catfish to be killed, and eradication of goldfish may be covered under regional council pest management plans such as in the Waikato Region of Aotearoa. Despite these latter attempts to ‘cover these gaps’, such omissions are worrying, especially given that the freshwater ornamental aquarium trade is arguably the greatest current biosecurity threat to freshwater ecosystems in Australasia (
The continuing tension between different legislation, governing organisations, and management objectives has been at least partially acknowledged. A recent proposal sought to create a new special permit to enable all species that can be defined as pest fish in areas where a specific problem has been identified, to be managed or eradicated under a single purpose, regardless of any other legislation (
The sustainability of the freshwater aquaculture sector depends on minimising the environmental impact generated by freshwater farms (
Fish farmer education and biosecurity initiatives do in their own way constitute negative socio-ecological feedback loops with respect to non-native fish (see Figs
In 1987, a proposal was made to introduce channel catfish (Ictalurus punctatus) from North America for aquaculture (
In 1989, a fish farm in the Auckland region was initially granted permission to farm the Australian Marron Crayfish Cherax tenuimanus at one location (
These cases demonstrate how the misalignment of legislation at the time allowed the legal importation of two INNS into Aotearoa despite their known histories of ecological damage in other jurisdictions, before subsequently legally preventing their release within the country. Both cases also demonstrate the continuing tension between economic interests and ecological protection as drivers in decision-making over the management of non-native fish (see
Although many translocations of freshwater non-native fish in Aotearoa happened in the past, due to the activities of acclimatisation societies, legislation now restricts translocations to native fish only (section 26ZM ‘Transfer or release of live aquatic life’ of the 1987 Conservation Act (2019 amendment). The current government approval process has two broad pathways: First, releasing species where they don’t already occur, which includes stocking a freshwater species at a fish farm for the first time, and second, releasing a species where it already occurs. Because only indigenous species are legally allowed to be translocated, a positive (reinforcing) feedback loop can be established, where native freshwater species, valued for conservation purposes and native fish farm expansion to supplement native populations under pressure or decline, drive more translocations of selected native species (Fig.
As discussed earlier, there are two notable NNS exceptions to the ’native only’ translocation policy and these are the grass and silver carp which are legally termed ‘restricted species’, as opposed to ‘pests’. Their translocation and release are allowed under licence provided they are ‘under control’. However, this system is not foolproof and there have been instances of previously legally released carp escaping to water bodies, other than the ones allowed in the licence (
Fish passage in Aotearoa is managed to prevent the passage of ‘undesirable’ fish species in order to protect ‘desirable’ fish species, their life stages, or their habitats’ (
As discussed previously, the acclimatization societies were dynamic drivers of fish introductions to bolster tourism and sport in the fledgling British colony of New Zealand (
Being the only country in the world where trout farming and commercial sale of trout is banned, government documents with any reference to trout culture use the term ‘trout production’ to distinguish it from all other freshwater ‘finfish aquaculture’. This is because although commercial farming is prohibited, hatcheries continue to produce trout for re-stocking purposes for recreational angling. The situation is culturally and politically complicated, with some iwi (Māori extended kinship group(s)) in some regions wanting the law changed to allow farming of trout (
Non-native fish introductions can drive policy-based socioeconomic positive feedback loops that increase fish transport out of source regions over time, as the perceived success and value of the introduced fish in recipient areas induces a ‘socio-economic invasion meltdown’, whereby economics drive further introductions as part of a positive feedback loop (Fig.
The debate over non-native freshwater fish management continues, and that is the ‘new reality in New Zealand’ (
Currently, the prevailing legislation, despite its shortcomings, seeks to promote negative feedbacks mitigating against the establishment and spread of INNS, and positive feedbacks that reinforce the recovery, persistence and growth of culturally important native species and fisheries. Continued use of language and selective ‘labelling’ of some INNS as ‘pests’, ‘noxious’ and ‘undesirable’ is a powerful tool in policy terms to promote negative feedbacks seeking to detect, manage and eliminate INNS. However, it should be acknowledged that some non-native fish such as brown trout are now so widespread in Aotearoa and have already drastically changed the trophic ecology of freshwater systems, that it is probably impossible to eradicate them, even if they are increasingly regarded as pests that need to be removed so native biodiversity can be restored (Chadderton 2001;
In Aotearoa, arguments over ecosystem restoration from a western fisheries perspective, with a status quo of established non-native sport or aquaculture species desirable to some stakeholders, as opposed to ‘reimagining’ fishery management with indigenous knowledge and values as a focal point, continue. Māori values and perspectives concerning freshwaters, which were all too often ignored in the past (
We would argue the current legislative landscape in Aotearoa is a patchwork, ‘make-do-and-mend’ approach to freshwater fish management, rather than a coordinated, coherent pragmatic approach. We would advocate the latter, until a much wider national debate has taken place on what people need and want from their freshwater fisheries in Aotearoa and how this could be achieved. A coordinated approach would require government organisations, iwi entities, and different stakeholder groups working together in a more equitable, unified way than previously witnessed. Only by doing this, can modernised, coherent policies and legislation be produced, whereby gaps, inconsistencies, anomalies and duplication can be minimised and when identified, resolved quickly. Using a SES approach with a focus on causal-loop relationships and feedbacks will be extremely valuable to improve understanding of the dynamics of drivers and outcomes underlying current non-native freshwater fish polices and how these can be manipulated to achieve agreed outcomes.
This research was conducted through the Fish Futures MBIE Endeavour programme, contract CAWX2101. Thanks to Joanne Clapcott for reviewing this manuscript. We thank both the editor and reviewer for their very constructive comments.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This study was funded by the Ministry of Buiness, Innovation and Employment, through the Fish Futures MBIE Endeavour programme.
Conceptualization: CM. Methodology: CM. Writing – original draft: CM. Writing – review and editing: JA, RH, EC, KM.
Calum MacNeil https://orcid.org/0000-0003-0402-9292
Robin Holmes https://orcid.org/0000-0002-8737-2717
Edward Challies https://orcid.org/0000-0003-0689-858X
Kiely McFarlane https://orcid.org/0000-0003-0321-2028
All of the data that support the findings of this study are available in the main text.