Outline of INVASAL White paper

Title2: Identifying knowledge gaps and key objectives associated with salmonid invasions in Chile

Authors’ list (proposal): Gomez-Uchida D, M Sepúlveda, S Neira, B Cid-Aguayo, B Ernst, TA Contador,

CB Canales-Aguirre, P Bahamonde, C Quezada-Romegialli, A Ramírez, JM Yáñez, LW Seeb, JE Seeb, I
Vila, TP Quinn, DE Schindler, BJ Hicks & C Harrod

Target journal: BioScience (IF: 5.4). Máx 5,000 words excluding refs, tables, figs, but includes textboxes.

Abstract

Chile is a long, geographically and socially diverse country, displaying strong latitudinal and altitudinal
gradients, and extreme ranges of climatic conditions, resulting in a wide variety of aquatic habitats. Such
habitats encompass the known native range for salmonid fishes, and likely has done much to support
their establishment and spread. Introduced salmonids have dramatically altered economic and social
landscapes in the region: to date, attempts to quantify their ecological impacts on native ecosystems and
taxa, and their contributions to the wider socio-economic system in the region have been dispersed. Here
we outline a cross-disciplinary approach to identify key knowledge gaps, indicative for other regions
experiencing or facing similar issues. Major gaps included the need to (1) characterise distribution of both
salmonid and native fishes, (2) evaluate ecological interactions between native taxa and salmonids and to
understand their impact at the ecosystem level, (3) understanding the societal importance of salmonid
fishes, and (4) the development of methods and policy for management and control of salmonids to
control strategies via suppression or eradication. We argue that management goals around salmonid
invasions are more likely to succeed if focused on both species and region-specific approaches,
recognising unique social, economic, and ecological settings. Changes in policy and education of society
at large about these invasive, yet valuable species, must hinge on cross-disciplinary research and a
network of scientists from diverse backgrounds.

Keywords: Salmonidae, fisheries, aquaculture, management, invasive species, social science

Human-mediated introductions of non-native species have profound consequences on native receiving

ecosystems. Non-native species that become invasive (i.e., establish, spread and negatively affect an
ecosystem) may cause community changes and population declines among native species, leading to
loss of ecosystem services. They can ecologically interact with multiple species and have far reaching
community consequences, acting as predators, prey, or both, as well as competitors and vectors of
pathogens, parasites or contaminants. Disentangling these roles is crucial to developing both the efficient
management of invasive species and effective conservation strategies for native species. However,
biological invasions are complex phenomena that also include multiple social dimensions. Invasive
species place us at the interface between both social (power-work-discourse) and natural realms,
extending the issue into the field of political ecology {Latour, 2004 #7045} . Therefore, invasive species
must be analysed considering specific impacts they have on the livelihoods, landscape, ecosystem
knowledge, mental, physical, social and psychological well-being of communities {Fitzgerald, 2009 #7041}
{Fortwrangler, 2013 #7042}. From a utilitarian perspective, not all invasive species appear harmful; some
may prime the development of emerging economies {Beever, 2019 #7047}. A participatory work
approach with local stakeholders is thus needed to tackle complex socio ecological phenomena {García-
Llorente, 2008 #7046}.
Salmonids (salmon and trout; family Salmonidae, Box 1) have been introduced around the globe owing to
their value as game and aquaculture fish. They have become successful invaders in both marine and
freshwater environments, although resident life-history forms have been in general more successful than
anadromous forms (Box 1; Quinn 2018). In South America, and Chile in particular, the presence of
salmonids is a controversial topic, because it carries both perceived negative and positive impacts.
Negative impacts are invariably associated to ecological interactions between salmonids and native taxa,
especially freshwater fishes. Positive impacts are invariably associated to employment, food provision
and recreation; salmonid fishing promotes turism and subsistence fishing for local communities free
capture from the scapees, , ocean nutrient subsidy, improve economy in local communities

Box 1: A short primer of salmon biology

Salmonids are a group of fish species native to the Northern Hemisphere, are important subjects in
ecology and evolution, and support recreational, commercial, and tribal fisheries. They show
significant variation in life history, including semelparous (one reproductive event) or iteroparous
reproduction (multiple reproductive events), anadromous (migrate to sea to feed but return to breed
in freshwater) or resident forms (spend entire life cycle in freshwater), and yearly or sub-yearly
residence of juveniles in freshwater before migrating to sea. They also home to their natal streams
with extraordinary precision to breed, which permits the evolution of reproductive isolation and local
adaptation between populations at various scales, ranging from tributaries to entire watersheds.
Salmonids are high-trophic level predators and can be considered keystone species especially in
relation to breeding. Nest digging by female salmon to deposit their eggs results in disturbance of the
riverbed affecting nutrient cycling and ecosystem metabolism. Carcasses from semelparous species
generate large amounts of marine-derived elements available for consumption and recycling by
resident species (both salmonids and non-related species), and elements are often transferred from
the river channel to riparian areas by floods and by predators and scavengers.

Specify what this paper is and is not – it is not another review, but rather a manifesto. Chile as a case
study.

The isolated and highly endemic south western region of South America threatened by invasive
salmonids

Here we focus on Chile as a microcosm of other regions around the globe that face the threat of
non-native salmonids and other invasive species. L ontinental
Continental Chile represents a microcosm of much of the globe: located in the south-western extreme of
South America, continental Chile it extends from the tropics to the sub-Antarctic. The country extends
across is characterised by , and marked latitudinal (17º—56ºS) and altitudinal gradients
(0 ~ 7000 m asl)
result ing in extreme ranges of climate conditions
that include arid, Mediterranean, temperate, high tundra, and polar climates (REF)
(Fig. 1).

Associated with this variation is the existence of diverse aquatic habitats that have been readily colonised by salmonid
Extreme latitudinal range of bioclimatic types have co-evolved during the Neogene (c.a. 23 Ma) along with
the final uplift of the Andes Cordillera (cita), and thus have generated short and steep flowing rivers
draining from the east to the west that nowadays are not connected to each other. In these particular
environments a unique and highly endemic ichthyofauna have evolved that is characterized as having
small sizes, retaining primitive characters and being adapted to rivers with high slope and variable stream
flow (cita). Jointly with fishes, entomofauna shows/have … . Salmonid fishes now dominate a wide range
of continental habitats including lakes, rivers and streams in lowland habitats, but also in extreme
altitudes (4500 m), in the driest desert in the world (River Loa), through to the most southern rivers in S
America.

Hotspot of biodiversity… endemic fishes… invertebrates…

How invasions by non-native salmonids have changed social and economic landscapes amid
scattered efforts for quantifying ecological impacts on native ecosystems

Salmonids were successfully introduced to Chile as a result of both aggressive government- and
privately-funded initiatives implemented since the start of the twentieth century. These included stocking
of lakes and river systems to promote recreational and commercial fisheries, including “sea-ranching”
initiatives among anadromous salmonids (1900 - 1990), and the rapid development of salmonid
aquaculture (1980 - present), both in freshwater and marine coastal areas. At least five species of
salmonids from Pacific (genus Oncorhynchus) and Atlantic lineages (genus Salmo) have established and
spread in Chile and South America (Table 1). The introduction and establishment of salmonids has been
a remarkable success, providing access to jobs, food, and recreation to regions with little economic
opportunities. Naturalized salmonids currently support recreational fisheries and emerging, small-scale
commercial fisheries. Salmon and trout aquaculture have positioned Chile as the world’s second largest
salmon producer, but at high environmental costs: habitat degradation, possible links to algal blooms, and
impacts of salmonid escapes from aquaculture facilities. Today, populations of non-native salmonids are
found in almost every major watershed, from the northernmost region and border with Peru, to Navarino
Island in the southernmost region of Magallanes province.

Understanding the impacts of salmonids at various levels of biological organization is necessary to

balance the importance of their presence in Chile’s ecosystems. However, salmonids are also highly
invasive and have extremely adverse impacts on native ecosystems. In Chile, the presence of salmonids
is controversial, and has marked implications for the socioeconomic and ecological well -being of the
country and the natural resources on which it depends. There is a pressing need to count with evidence-
based policy for the future management of invasive salmonids in Chile by asking What and where are
the ecological and socioeconomic impacts of invasive salmonids in Chile and how this
information can be used to develop and implement evidence-based management solutions and
(through?) policy?

The mission of the Millennium Nucleus of Invasive Salmonids (INVASAL) is to: (1) undertake world-
class research with national and global impact using cutting-edge techniques to characterize the
distribution and impacts of free-living salmonids on native ecosystems, both in fresh- (lakes rivers and
streams) and coastal-waters, including population-, community- and ecosystem-level effects, and their
societal impacts; (2) understand how Chilean society uses and values free-living salmonids; (3) develop
tools for resource managers, conservation biologists, enforcement personnel and industry and
disseminate findings to them and other stakeholders including fishers, NGOs and society at large, in order
to improve and develop efficient policy and management strategies for both invasive salmonids and
native aquatic communities; and (4) mentor students, professionals, and young researchers in a friendly,
welcoming environment where equality is strived for.

Introduced salmonids have dramatically altered economic and social landscapes across the world, and
particularly so in Chile. Although individual attempts have been made to quantify the ecological impacts of
salmonids on native ecosystems and taxa, and their contributions to the wider socio-economic system,
efforts have dispersed. There is now a recognition that a more coherent, multidisciplinary approach is
required to direct future management. However prior to providing suggestions regarding a roadmap for
future activities, it is necessary to identify the gaps in our knowledge so that they can be bridged (Chen et
al., 2017).

Here we outline key knowledge gaps that limit our capacity to develop science-based policy to manage
ecological and societal impacts of non-native salmonids in Chile. Although we have focused on Chile,
given the geographical, social and ecological diversity of the country, our observations likely extend to
other regions in the southern hemisphere (and beyond) already experiencing or facing the future spread
of non-native salmonids and other taxa.

Knowledge gap #1: Mapping the current distribution and abundance of salmonids & native fishes

There is a pressing need for current information on their status, their actual - not perceived - impacts on
receiving habitats, native species, and supporting ecosystems and their importance to and use by human
populations. Ddistribution and impacts of free-living salmonids on native ecosystems, both in fresh- (lakes
rivers and streams) and coastal-waters. This will likely require a combination of field-based sampling,
interviews from several stakeholders (e.g., fisheries scientists, anglers, fishing guides, hatchery managers
and others stakeholders), literature review and even social media monitoring should yield the most
comprehensive data set to date. These data will be integrated using a geographic information system into
a dynamic and open access map of the distribution of salmonid and native fishes in Chile.

We will use the cutting edge approach of the analysis of environmental DNA (eDNA) for rapid assessment
of the presence of native and non-native fishes in rivers across continental Chile. This involves filtering
water from a given habitat and extracting DNA to identify species composition present at a site using both
metabarcoding (i.e. next-generation sequencing of entire communities of fishes) and real-time quantitative
PCR (i.e. highly sensitive detection of traces of species-specific DNA present in the water).
Currently the use of eDNA is revolutionizing the assessment of fish distributions and by undertaking
national-scale study…

We will also use state-of-the-art genomic techniques, including high-throughput genotyping gentoyping
and next-generation sequencing, to assess population genetic patterns in order to better understand past
and present demographic processes experienced by invasive salmonid populations. In this regard, our
primary interest will be focused on the study of natural study natural selection and local adaptation events
and their effect on shaping the genome of invasive salmon populations.
We expect that these approaches will provide further insights into the biological basis of their i

Knowledge gap #2: Evaluating ecological interactions between native taxa and salmonids and
their ecosystem role
The biomass of salmonids in Chilean freshwaters is likely greater than for all native fishes combined,
reflecting their innate plasticity and flexible diet, and potential competitive superiority over native fishes,
which in freshwater tend to be endemic, small bodied and less aggressive. Salmonids can attain large
individual sizes, consuming increasing amounts of invertebrates and fish as they grow. In marine habitats,
Chinook salmon can gain ca. 10kg a year feeding on small fish and krill. Conversely, despite undergoing
predator release in Chile, invasive salmonids represent an ecological opportunity for piscivorous
predators, both native (e.g. marine mammals, fish eating birds) and introduced (American mink), and
predation on salmonids has been markedly understudied in Chile.

Contamination is known to be a direct driver of ecosystem change at the global scale, contributing the loss of biodiversity and degrada

Few studies reported increasing POPs concentration during the last 30 years in remote lakes located in Patagonia, by using sedimentary

s Salmon feed largely on small pelagic fish such anchovy (Engraulis ringens), common sardine
(Strangomera bentincki) and Falkland sprat (Sprattus fueg uensis). This interaction may have important
consequences for ecosystem-based management approaches, since the latter species support important
fisheries and also play a critical ecological role by channeling plankton productivity towards higher trophic
levels (Cury et al., 2000). Energy subsidies and ecosystem implications of Chinook s Salmon in the
coastal ocean and in many watersheds that drain to the Pacific Ocean are currently unknown, and
knowledge of how marine-derived nutrients in freshwater introduced by semelparous salmonids (see Box
1) have changed the water-land interphase in upper river systems is lacking. It is possible that invasive
Chinook salmon are facilitating invasions and establishment of non-native rRainbow tTrout, bBrown
tTrout, or both, that capitalize on the marine subsidy provided by spawning fish. In addition to being a top-
level predator, Chinook s Salmon have become a significant prey item among marine mammal
populations such as sea lions, suggesting potential positive impacts on mammal populations that need to
be quantified. Chinook s Salmon have also become a significant prey item for some marine mammals and
other predators such as sharks along their native range (e.g. {, #6540@@hidden;,
#6538@@hidden}Adams et al. 2016, Chasco et al. 2017, Seitz et al. 2019). For instance, an increase in
the abundance of pinniped (seals and sea lions) and killer whale species, has reduced Chinook salmon
abundance in the northeastern Pacific Ocean {Chasco, 2017 #6538}. Outside their native range, New
Zealand fur seals (Arctocephalus forsteri) in Marlborough Sounds, New Zealand, consume farmed
Chinook salmon {Kemper, 2003 #7040}, and of free-living Chinook salmon by South American sea lions
(Otaria flavescens) in Toltén River, in southern Chile {Gomez-Uchida, 2016 #2345}, which suggest,
potential positive impacts of the presence of this non-native fish on these marine mammals. An estimate
of the consumption by these and other potential predators is needed to improve our understanding
predator-salmon relationships and to inform fishery management decisions.

In addition, the presence of animal migrations is responsible for the transport of nutrients, energy and even pollutants across different e

Few studies reported increasing POPs concentration during the last 30 years in remote lakes located in Patagonia, by using sedimentary

have the have the potential to affect native taxa and ecosystems across multiple levels of biological and
socio-ecological organisation. With this in mind there is a need to follow a multidisciplinary approach to
examine ecological interactions between salmonids and native species that extends from primary
producers, through to putative prey, competitors, and predators.

Salmonids life cycles characteristically include spawning migrations that lead to individuals moving across ecosystem boundaries inclu

Salmonids Salmonid predation over zooplankton may trigger a phytoplankton bloom…

We will characterize the trophic ecology of species and functional groups through an innovative approach,
combining the use of gut contents, stable isotope analysis, gut content DNA-barcoding, and
bioenergetics/food web modeling.
We will study this at a series of locations indicative of Chile’s climatic zones and the diversity of
management and ecological challenges associated with salmonid fishes in both freshwater and marine
habitats (Fig. 2). Since 2013, the Chilean Fisheries and Aquaculture Law includes the ecosystem
approach to management as a way of securing sustainability of stocks, commercial activities, and the
safeguard of ecosystems. Therefore, we will develop qualitative and quantitative ecosystem models that
will allow us to synthesizes ecosystem and socio-economic impacts of non-native salmonids on Chile’s
aquatic environments. Later, these models will inform managers on the likely consequences of different
management measures (e.g., control, eradication, exploitation) applied to salmonids.

The presence of salmonids potentially influences life history traits of key native fishes and invertebrates
as well as ecosystem functioning.

Knowledge gap #3: Understanding the importance of salmonids for Chile’s society

Chile is globally known for the recreational fisheries targeting naturalized salmonids (Arismendi &
Nahuelhual 2007). However, there is also illegal, unregulated and unreported (IUU) artisanal small-scale
fisheries of salmon populations, e.g. Chinook salmon, providing important economic support to low-
income communities (Musleh 2016). In response to this situation, an emergent governance and
management strategies for these fisheries is being advanced by fisheries authorities and concerned
stakeholders (DS 124/2018 MINECON, Gomez-Uchida et al. 2016). In addition, Chile has implemented in
2013 changes in Fisheries law whose core objectives are the conservation and sustainable use of its
living marine resources. This legislation mandates scientifically based fisheries management using
precautionary and ecosystem approaches (Ley de Pesca y Acuicultura n°20.567, 2013).
This new legislation creates a n window of opportunity to guide this emergent governance, towards
implementation of elements of the ecosystem approach to fisheries management, in small-scale fisheries
targeting non-native salmonids.

including recreational and commercial opportunities,

To better meet these challenges, researchers and practitioners focus on understanding linkages between
social and ecological dynamics — often referred to as linked or coupled social-ecological systems and
how these dynamics affect the fisheries sustainability (Degnbol & Mc Kay 2007). This is part of a more
comprehensive view for the implementation of the ecosystem-based fisheries management approach
(Kittinger et al. 2013). Emerging governance towards ecosystem-based management in non-native
salmonid fisheries requires an integrated and participatory social-ecological approach to develop
implementation frameworks and management strategies (figure ) (Pascual et al. 2009, Fulton et al. 2014,
Folke et al.2016). i) analyse the structure and dynamics of the social-ecological systems involved in the
small scale fisheries targeting non –native salmonids in Chile; ii) reveal components and processes
relevant to the social-ecological fit of emerging governance and management; and iii) understand the
potential sustainability outcomes of emerging governance and management strategies in the small-scale
salmonids fisheries.
To achieve this, we will implement a participatory social-ecological system analysis framework (figure),
involving key stakeholders in order to develop SES models, build desired governance regime scenarios
and test management strategies derived from those scenarios. (Fulton et al. 2015, Ortiz and Levins 2011;
Walker et al. 2006). From a stakeholder viewpoint, this approach focus on a participative process of
building a shared understanding, of the elements that generate and modify the capacity of a fisheries
social-ecological system, to deliver environmental benefits (catch, recreation, traditions, etc) contributing
to desired levels of social wellbeing (Walker et al. 2002, Hillborn et al. 2015, Fletcher et al. 2014).
Chile is globally famous for the quality of its recreational fisheries targeting naturalized salmonids.
However, there is also small-scale exploitation of abundant anadromous salmon populations, e.g.
Chinook salmon. They provide major economic support to low-income communities. However, there are
potential issues regarding contamination (as upper trophic level predators) e.g. with metals and this is
largely unknown at the moment. For conservation and management policies to succeed, different
stakeholder values must be considered as salmonid fishes mean different things to different stakeholders.
Salmonids represent an essential source of income to some; to others, they are a form of biological
contamination to be removed to protect native species and ecosystems.

Using interviews and regional workshop-based discussions, we will assess public and stakeholder opinion on the value of salmonid and

.
The Lake and Aysén Districts of Chile support the bulk of national aquaculture activity. Aquaculture is key
economic concern in Chile, providing >70% of export income from the south of Chile and employment to
many people. It continues to expand into new locations, representing a major source of potentially
invasive fish wherever fish are cultured. Three commercially important salmonid species unintentionally
escape from aquaculture facilities: Atlantic salmon, coho salmon and rainbow trout. INVASAL researchers
showed that naturalized rainbow trout were genetically divergent from cultured fish. Cultured and
naturalized salmonids are isotopically distinct too, and we will develop genetic-isotopic tools to identify
whether fish are cultured, naturalized or escapees from aquaculture. This will be important to fully
understand the relative ecological impacts of invasive salmonids on marine ecosystems associated with
naturalized and recent escapees.

Given the scale of the potential resource, and the existence of unregulated fisheries for naturalized and
aquaculture-escaped salmon, there is a large potential market for unofficial salmonid products, likely sold
under the label of Atlantic salmon (the most farmed species in Chile). We will examine the use of
naturalized and escapee salmonids in the human food chain after identifying tissues to species by DNA
barcoding and origin (farmed vs. free-living) by stable isotope analysis.

INVASAL will also continue providing estimates of abundance of returning Chinook salmon by using field-
methods to measure abundance (hydroacoustics, spawner and redd surveys) and catch data provided by
stakeholders. These data will allow us to characterize the scale and location of the problem.

Recreational fisheries and farming of non-native salmonids in Chile have been successful, generating
access to jobs, food and recreation (Pascual et al. 2009). However, salmonids are highly invasive with
negative impacts on native ecosystems with associated economic costs (Sepúlveda et al. 2013,
Arismendi et al. 2014, Penaluna et al. 2009). This creates a challenging conservation and management
trade-off between social benefits derived from fishing salmonids and the negative impacts on native
biodiversity.

Knowledge gap #4: Developing methods & policies for the management & control of invasive
salmonids
In Chile, salmonids have seized in the last century an ample variety of aquatic systems, including an
unaccounted number of rivers, lakes and creeks from the northern desert to Cape Horn and from the
coast to the Andes. This vast geographic range extends over private and public property, including
national parks and involves a great number of stakeholders. In the absence of a national unified stance
on the role of free-living salmonids in the country, legislation has not evolved accordingly to neither
safeguard native fauna from negative biological interaction nor to assure the provision of technical
support to adopt properly designed management actions to control or eradicate these species.
Controlling invasive species is difficult, but some INVASAL workers have led successful efforts to remove
invasive species to conserve native species. We will undertake an effort to remove rainbow trout from the
Chungará system in northern Chile. Lake Chungará, one of the most important freshwater ecosystems in
the Chilean Altiplano, supports a single native species, the endemic Orestias chungarensis. However,
rainbow trout were illegally introduced during the mid-20th century and potentially compete with, and prey
on O. chungarensis. INVASAL scientists are undertaking innovative research to assess the status and
identify biological bottlenecks of this invasive population for designing a proper cost/effective eradication
approach. This flagship program acts as a case study and will provide guidance for similar activities in
Chile and worldwide.
Sizing biological and socioeconomic impacts of free-living anadromous salmon is another topic addressed
with great vigour by INVASAL scientists. We provided scientific and management support to the Chilean
National Fisheries Authority on developing the first fisheries decree to regulate artisanal fishing on
Chinook salmon in Toltén estuary. This case study comprises a great socioecological model for dozens of
watersheds in southern Patagonia. We will estimate the population size of the rainbow trout population
using a series of methods in the lake and inlet river, providing an indication of the number of fish to be
removed. Adult rainbow trout spawn in the inlet Chungara River and will be trapped and removed during
their spawning migration through a single fixed point. Juvenile fish will be removed after being trapped
and electrofished. Visual surveys will be used to identify and destroy eggs buried in the river bed. We will
continue monitoring trout abundance throughout the project and feel that it is feasible that in a 3-year
programme that a major rainbow trout population can be reduced to a point where it is no longer self-
sustaining. This flagship program act as a case study and provide guidance for similar activities in Chile
and worldwide.

Education and outreach

This cross-cutting theme joins the four data-driven themes. The INVASAL team incorporates professional
educators and scientists strongly involved in education and outreach across four continents, and the
project will inform and train the next generation of scientists and citizens. Students and early career
scientists will undertake practical training in the various laboratories in Chile and in our partner
organizations in the US and NZ and annual summer schools. We will inform the next generation of
scientists about the issues associated with INVASAL through school visits and field-schools organized
across the different regions by the excellent CONICYT PAR-EXPLORA scheme.

Several members of the INVASAL team have strong social media presences both in Chile and
internationally. We will maximize publicity and interactions with the public through a dedicated web page
linked to social media accounts detailing the issues behind INVASAL, our activities and production of key
milestones. We will strengthen existing relationships with the traditional media across the regions of Chile,
allowing wide geographical dissemination and the decentralization of science news, which in Chile, is
typically Santiago-centered, unlike the issue of invasive species.
Education for citizens: In Chile, and maybe in other countries, local citizens think that salmonids are native of their own country. Also

Box 2: Rainbow trout in Lake Chungara – an immediate threat to Orestias chungarensis

There is no documentation on how rainbow trout were illegally released into Lake Chungará.
Anecdotical information suggests that multiple introductions may have taken place. Captive
broodstock may have been released following a failed attempt of aquaculture in tanks, anglers may
have stocked the lake to naturalize the species, or both. Rainbow trout evolved a migratory life
history: adults feed in the lake, preying on endemic karachi O. chungarensis as well as benthic
amphipods, but spawn at “bofedales”, a network of inlet streams spreading over XX km2 that
comprise Chungará River. Juveniles can remain up to ? years in the river.

{, #7038@@hidden}
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Table 1. Most important non-native salmonids naturalized in South America and Chile and perceived
relative ecological attributes and socioeconomic value.

Species Life history Latitudinal Abundance Aquaculture Recreational Co

distribution Fishing
Chinook salmon Anadromous, Very Wide (+++ Very High (++ Very Low (+) Very High (+ Ver
O. tshawytscha Semelparous +) +) +++)
Fifty S
watersheds from exp
Imperial River the
(Pacific Ocean) To
to (Santa Cruz
River Atlantic
Ocean

Resident, Very Wide (+++

Iteroparous +)
Over a hundred
No
Rainbow trout watersheds from Very high (++
+++ ++++
O. mykiss Chungará River ++)
to Navarino
Island (Pacific
Ocean)

Coho salmon Anadromous, Narrow (++) Low (++) ++ High (+++) No

O. kisutch Semelparous

Cherry salmon Very Narrow (+) Unknown (o) No

O. masou
Atlantic salmon Very Narrow (+) Unknown (o) Very Important + No
S. salar (++++)
Brown trout Wide (+++) Unknown (o) o +++ No
S. trutta
Brook charr Very Narrow (+) ++ o ++ No
S. fontinalis
References: 1.{Arismendi, 2007, Non-native salmon and trout recreational fishing in Lake Llanquihue`,
Southern Chile: Economic benefits and management implications}; 2{Arismendi, 2009, Aquaculture`, non-
native salmonid invasions and associated declines of native fishes in Northern Patagonian lakes};
3{Arismendi, 2011, Seasonal age distributions and maturity stage in a naturalized rainbow trout
(Oncorhynchus mykiss Walbaum) population in southern Chile reveal an ad-fluvial life history}; 4{Astorga,
2008, Naturalized Chinook salmon in the northern Chilean Patagonia: Do they originate from salmon
farming?}; 5{Correa, 2008, Chinook salmon invade southern South America}; 6{Faundez, 1997, Allozyme
variability in brown trout Salmo trutta in Chile}; 7{Habit, 2010, Changes in the distribution of native fishes
in response to introduced species and other anthropogenic effects}; 8{Niklitschek, 2011, INFORME
FINAL PROYECTO FIP 2008-30 “EVALUACIÓN CUANTITATIVA DEL ESTADO TRÓFICO
DESALMÓNIDOS DE VIDA LIBRE EN EL FIORDO AYSÉN`, XI REGIÓN”}; 9{Soto, 2001, Escaped
salmon in the inner seas`, southern Chile: Facing ecological and social conflicts}; 10{Soto, 2006,
Southern Chile`, trout and salmon country: invasion patterns and threats for native species}; 11{Soto,
2007, Establishment of Chinook salmon (Oncorhynchus tshawytscha) in Pacific basins of southern South
America and its potential ecosystem implications}.
Figure 1. Summary of ecological and social settings relevant to study impacts of the presence of invasive
salmonids on ecosystems and society.
Stomach content analyses suggest that in their marine phase, Chinook Invasive salmonids