Species movements within biogeographic regions: exploring the distribution of transplanted mollusc species in South America

Abstract

The movement of species is among the most serious environmental threats of the new millennium, as the transplantation of species beyond their native or historical range has intensified in the last five decades. Traditionally, studies on bioinvasions have focused on species that have been introduced, deliberately or accidentally, to biogeographic regions where they did not previously occur. However, less attention has been given to species movement to novel areas within the same biogeographic region. Our research group, the South America Introduced Molluscs Specialists, analyzed potential cases of native South American mollusc species introduced deliberately or accidentally beyond their natural range within South America. To achieve this, it is key to differentiate between anthropogenic processes and passive responses to environmental conditions. We considered the past and current spatial distribution of species, analyzed known or putative vectors, and discuss the impacts of taxonomic and nomenclatural knowledge. Based on the evidence currently available, we propose different scenarios to explain observed changes in mollusc distributions within South America. Seventeen transplanted mollusc species (TMS) were recognized, considering marine, freshwater, and terrestrial environments. Of the 189 South American ecoregions 31 were occupied by transplanted species, but this proportion varied by environment: 10 of 28 marine ecoregions, 12 of 52 freshwater ecoregions, and 9 of 109 terrestrial ecoregions. The ecoregions with TMS are generally located in the peripheral zones of the continent. Those with the highest number of TMS were the Southern Caribbean (three species) in the marine environment, the Central Andean Pacific Slopes (three species) in the freshwater environment, and the Alto Paraná Atlantic forests (two species) in the terrestrial environment. The number of unintentionally moved TMS is greater than those moved intentionally. The transplantation process is similar to the introduction and settlement process of non-native mollusc species, and so is their impact.

Full text

Vol.: (0123456789)
1 3
Biol Invasions
https://doi.org/10.1007/s10530-022-02942-z
REVIEW
Species movements withinbiogeographic regions: exploring
thedistribution oftransplanted mollusc species inSouth
America
GustavoDarrigran · IgnacioAgudo‑Padrón · PedroBaez· CarlosBelz · FranzCardoso ·
GonzaloA.Collado · ModestoCorreoso · MaríaGabrielaCuezzo · CristinaDamborenea ·
AlejandraA.Fabres · MonicaA.Fernandez · SuzeteR.Gomes · DiegoE.GutiérrezGregoric ·
SergioLetelier· CésarLodeiros · SandraLudwig · MariaCristinaMansur·
SamuelNarciso· GuidoPastorino · PabloE.Penchaszadeh· AnaCarolinaPeralta ·
AndreaRebolledo· AlejandraRumi· RodrigoB.Salvador · SoniaSantos · PaulaSpotorno ·
SilvanaCarvalhoThiengo · TeofâniaVidigal · AlvarCarranza
Received: 9 September 2021 / Accepted: 1 October 2022
© The Author(s), under exclusive licence to Springer Nature Switzerland AG 2022
have focused on species that have been introduced,
deliberately or accidentally, to biogeographic regions
where they did not previously occur. However, less
attention has been given to species movement to novel
areas within the same biogeographic region. Our
research group, the South America Introduced Mol-
luscs Specialists, analyzed potential cases of native
South American mollusc species introduced deliber-
ately or accidentally beyond their natural range within
Abstract The movement of species is among the
most serious environmental threats of the new millen-
nium, as the transplantation of species beyond their
native or historical range has intensified in the last
five decades. Traditionally, studies on bioinvasions
Supplementary Information The online version
contains supplementary material available at https:// doi.
org/ 10. 1007/ s10530- 022- 02942-z.
G.Darrigran· C.Damborenea(*)·
D.E.GutiérrezGregoric· A.Rumi
División Zoología Invertebrados, Museo de La Plata,
FCNyM, UNLP; CONICET, LaPlata, Argentina
e-mail: cdambor@fcnym.unlp.edu.ar
I.Agudo-Padrón
Projeto “Avulsos Malacológicos”, Florianópolis, SC,
Brazil
P.Baez· G.A.Collado· A.A.Fabres· S.Letelier·
A.Rebolledo
Sociedad Malacológica Chile, SMACH, Santiago, Chile
P.Baez
Centro de Investigación Marina de Quintay (CIMARQ),
Facultad de Ciencias de la Vida, Universidad Andrés
Bello, Quintay, Chile
C.Belz
Laboratório Ecologia Aplicada e Bioinvasões, Centro
Estudos doMar/Universidade Federal doParaná,
PontaldoParaná, Paraná, Brazil
F.Cardoso
Laboratorio de Biología y Sistemática de Invertebrados
Marinos, Departamento Académico de Zoología, Facultad
de Ciencias Biológicas, Universidad Nacional Mayor de
San Marcos, Lima, Peru
G.A.Collado
Departamento de Ciencias Básicas, Facultad de Ciencias,
Universidad del Bío-Bío, Chillán, Chile
M.Correoso
Departamento de Ciencias de la Vida y Departamento de
Ciencias de la Tierra y de la Construcción, Universidad
de las Fuerzas Armadas, ESPE, Valle de los Chillos,
Sangolquí, Ecuador
M.G.Cuezzo
Instituto de Biodiversidad Neotropical, CONICET/
Facultad de Ciencias Naturales UNT, Tucumán, Argentina

G.Darrigran et al.
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South America. To achieve this, it is key to differen-
tiate between anthropogenic processes and passive
responses to environmental conditions. We consid-
ered the past and current spatial distribution of spe-
cies, analyzed known or putative vectors, and discuss
the impacts of taxonomic and nomenclatural knowl-
edge. Based on the evidence currently available,
we propose different scenarios to explain observed
changes in mollusc distributions within South Amer-
ica. Seventeen transplanted mollusc species (TMS)
were recognized, considering marine, freshwater, and
terrestrial environments. Of the 189 South American
ecoregions 31 were occupied by transplanted species,
but this proportion varied by environment: 10 of 28
marine ecoregions, 12 of 52 freshwater ecoregions,
and 9 of 109 terrestrial ecoregions. The ecoregions
with TMS are generally located in the peripheral
zones of the continent. Those with the highest num-
ber of TMS were the Southern Caribbean (three spe-
cies) in the marine environment, the Central Andean
Pacific Slopes (three species) in the freshwater envi-
ronment, and the Alto Paraná Atlantic forests (two
species) in the terrestrial environment. The number
of unintentionally moved TMS is greater than those
moved intentionally. The transplantation process is
similar to the introduction and settlement process of
non-native mollusc species, and so is their impact.
Keywords Transfer· Translocation· Range
expansion· Bioinvasion· Ecoregions
Introduction
To develop control or eradication measures, the Con-
vention on Biological Diversity (CBD 1992, 2010)
prioritizes the identification and study of invasive
non-native species, one of the most significant causes
of biodiversity loss at a global scale (e.g., Blackburn
etal. 2020). This framework led specialists on mol-
luscs and bioinvasions from different regions of South
America to establish an expert research group, namely
A.A.Fabres
Laboratorio de Genética y Evolución, Departamento de
Ciencias Ecológicas, Facultad de Ciencias, Universidad de
Chile, Santiago, Chile
M.A.Fernandez· S.R.Gomes· S.CarvalhoThiengo
Laboratório de Referencia Nacional para
Esquistossomose-Malacologia, Instituto Oswaldo Cruz,
Fiocruz, RiodeJaneiro, Brazil
C.Lodeiros
Grupo de Investigación en Biología y Cultivo de
Moluscos, Dpto. de Acuicultura, Pesca y Recursos
Naturales Renovables, Facultad de Ciencias Veterinarias,
Universidad Técnica de Manabí, Portoviejo, Ecuador
C.Lodeiros
Instituto Oceanográfico de Venezuela, Universidad de
Oriente, Cumaná, Venezuela
S.Ludwig
Pós-Graduação em Genética, Inst. Ciências Biológicas,
Universidade Federal Minas Gerais, BeloHorizonte,
Brazil
M.C.Mansur
Grupo Pesquisa CNPq, Biodiversidade de Moluscos
Continentais, Museu de Ciências Naturais, PortoAlegre,
Brazil
S.Narciso
CIAC-FUDENA, Centro de Investigación y Atención
Comunitaria, Chichiriviche, Falcón, Venezuela
G.Pastorino· P.E.Penchaszadeh
Museo Argentino de Ciencias Naturales
“Bernardino Rivadavia”, CONICET,
CiudadAutónomadeBuenosAires, Argentina
A.C.Peralta
Laboratorio de Biología Marina, Departamento de
Estudios Ambientales, Universidad Simón Bolívar,
Caracas, Venezuela
R.B.Salvador
Museum ofNew Zealand Te Papa Tongarewa, Wellington,
NewZealand
S.Santos
Departamento de Zoologia, Universidade doEstado doRio
de Janeiro (UERJ), RiodeJaneiro, RJ, Brazil
P.Spotorno
Programa de Pós-Graduação em Oceanologia, Instituto
de Oceanografia, Universidade Federal doRio Grande,
FURG,, RioGrande, RS, Brazil
T.Vidigal
Laboratório de Malacologia e Sistemática Molecular,
Departamento de Zoologia, Instituto de Ciências
Biológicas; Lelf: Laboratório de Estudos de Limnoperna
fortunei, Centro de Pesquisas Hidráulicas, Universidade
Federal de Minas Gerais, BeloHorizonte, Brazil
A.Carranza
Área Biodiversidad y Conservación, Museo Nacional de
Historia Natural, 25 de mayo 582, Montevideo, Uruguay

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the “South America Introduced Molluscs Specialists
Group” (eMIAS; https:// alien mollu scsgr oup. wixsi te.
com/ saamg roup). Once established, an initial synthe-
sis of the status and distribution of non-native mol-
luscs in South America at an ecoregional scale was
developed (Darrigran etal. 2020). This work targeted
introduced species in South America from other con-
tinents. However, little attention has been paid to the
phenomenon of species movement within the same
biogeographic regions. In this context, eMIAS identi-
fied species of molluscs, native to South America, that
had experienced modifications to their historical dis-
tributions within the continent that are not attributable
to natural dispersal processes. Cases analyzed ranged
from obscure local species, such as the sea slug Aply-
sia dactylomela Rang, 1828, to globally conspicuous
species, such as the black-striped mussel, Mytilopsis
sallei (Récluz, 1849), a brackish-water bivalve native
to coastal lagoons and estuaries of Central America
and the Caribbean (Fernandes etal. 2018).
In the mainstream literature, the terms ‘trans-
plantation’, ‘transfer’, or ‘translocation’ have been
used interchangeably to identify this process. Even
though the terms are typically considered synony-
mous, several authors have pointed out important
distinctions and alternative definitions. Currently, it
has been argued that the planet is going through a
new era, the Anthropocene, in which disturbances
are mainly generated from the globalization of trade
and climate change (Hobbs etal. 2018). These pro-
cesses are advancing at unprecedented rates and
scales, and have yet to reach their climax. Because
of climatic changes, species can access new regions,
while others are forced to reduce their ranges (or
disappear) in response to altered environments.
Hayward etal. (2015) even proposed redefining the
IUCN guidelines on the natural range of species to
include potential range shifts and subpopulations
existing outside the natural range but in broadly
similar habitat. In this work, we follow the defini-
tion of the IUCN (2022), in which an indigenous
species (= native species) is defined based on three
criteria: (1) it is intrinsically part of the ecosys-
tem and developed there, (2) it arrived in the area
long before records of such matters were kept, (3)
it arrived by natural means (unaided by human
action). We acknowledge that this definition is only
a pragmatic and conventional rule to be used to
guide our study (Holguín 2003). Herein, a species
is considered non-native when it is introduced out-
side its natural geographical range through human
action, being able to maintain a self-sustaining
population (Turbelin et al. 2017). If a non-native
species disperses and has an evident environmental
and socio-economic impact, it is considered inva-
sive (Darrigran etal. 2020). Furthermore, we define
transplantation (transfer, translocation) as the move-
ment of native species from one locality to another
by human action, with successful establishment
outside their known historical geographical range
(in our case, within South America). Such species
movements can be both accidental or deliberate
(e.g., for conservation, trade, research). A species
is considered to be established in the environment
when there are populations with multi-generational
reproductive success (Kočovský etal. 2018).
Knowledge of the invertebrate fauna of South
America is still fragmented and incomplete (Dam-
borenea etal. 2020; de Barros et al. 2020). Despite
the high richness of native molluscs in the continent,
there are major knowledge gaps, mostly because of
the biased and unequal coverage by the areas studied,
errors in identification, and taxonomic uncertainty,
all of which can have consequences for conservation
(Salvador 2019; de Barros etal. 2020). There are also
differences among marine, freshwater and terrestrial
environments in terms of species richness, research
effort, and vulnerability. In contrast, the socio-eco-
nomic and ecological impacts of some invasive spe-
cies have led to a more comprehensive knowledge of
these species (Darrigran etal. 2020). As a corollary,
in South America, the distribution and biology of
economically important invasive mollusc species are
better understood than the distribution and biology of
native molluscs (Dos Santos etal. 2021). Similarly,
some of the knowledge of certain species native to
South America has come from studies on other con-
tinents where they have become invasive (e.g., Wong
et al. 2011; Joshi etal. 2017; Calazans et al. 2018,
2021; Fernandes etal. 2018).
In this context, this work aims to assess native spe-
cies of molluscs from South America that have been
transplanted within the continent, including marine,
freshwater, and terrestrial realms. Based on the evi-
dence currently available, we propose different sce-
narios to explain observed changes in molluscan dis-
tribution within the continent.

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Material andmethods
A collaborative effort was made by 29 expert mala-
cologists and taxonomists from countries in South
America (Argentina, Brazil, Chile, Ecuador, Peru,
Uruguay, and Venezuela) and from New Zealand,
belonging to the eMIAS group. Scientific and grey
literature, databases on the subject were analyzed,
and experiences were exchanged through a virtual
forum among the group. Then, the group listed the
transplanted mollusc species (TMS) and reached a
consensus on the status of each species in the 189
ecoregions (marine, freshwater, and terrestrial) recog-
nized in South America (Online Resources 1–3).
The geographic range of each species was deter-
mined based on literature records and expert experi-
ence at an ecoregional scale (see also Darrigran etal.
2020). Following Olson and Dinerstein (2002), ecore-
gions were defined as areas of land or water with a
characteristic set of natural communities, ecological
dynamics, and environment that share most of their
species. Geographic Information System (GIS) lay-
ers were based on Olson etal. (2001) for terrestrial
ecoregions, Spalding etal. (2007) for marine ecore-
gions, and Abell etal. (2008) for freshwater ecore-
gions. In South America, 28 ecoregions are recog-
nized for marine environments (http:// maps. tnc. org/
files/ metad ata/ MEOW. xml; Online Resource 1), 52
for freshwater environments (http:// maps. tnc. org/ files/
metad ata/ FEOW. xml; Online Resource 2), and 109
for terrestrial environments (http:// maps. tnc. org/ files/
metad ata/ TerrE cos. xml; Online Resource 3).
The criterion for a species to be considered as a
TMS was the existence of documentation of its trans-
plantation within South America. To avoid confu-
sion between the type of species under study and in
agreement with the definitions mentioned above, and
the description of introduction pathways by Faulkner
et al. (2020), we worked on the basis of three
premises:
1. The way in which the species are distributed,
both in geographic space and in time, responds
to historical, ecological, and physiological factors
throughout their range (Maciel-Mata etal. 2015).
2. The causes of transplantation of native species
are human-related, including conservation, trade,
research, and accidental transport.
3. Transplanted species can cause the same prob-
lems and potential dangers as introduced species,
which leads us to consider the same pathway(s)
as those proposed for introduced organisms by
the Convention on Biological Diversity (CBD
2014) (as interpreted by IUCN, 2017). The mech-
anisms and pathways by which TMS may arrive,
and enter a new region are detailed in Fig.1.
Based on these premises, the list of transplanted
species was compiled considering known histori-
cal distribution, new location(s), ecoregion(s) of the
new location(s), pathway(s) of transplantation, and
impacts. References are given for each case. Further-
more, mollusc species transported by humans and
only found in captivity were mentioned, but are not
Fig. 1 Classification frame-
work for routes of introduc-
tion / transplantation, modi-
fied from Faulkner etal.
(2020). The nomenclature
proposed in IUCN (2017)
was partially applied. The
mechanisms and pathway
categories are based on
Hulme etal. (2008)
Transplant
mechanism
Transplant
pathway
category
Transplant pathway
sub-category
Increasing human role
Moveme nt of
commodity
Ve ctor
Spread
Release
Escape
Contaminant
Stowaway
Corridor
A-Biological contro l; B -Conservation in wild ;
-Otherrele ase
A- Agriculture; B- Aquacult ure; C-Hortic ulture;
D- Ornamental; E-Otherescape
A- Parasite of animals; B- Parasite of plants;
C-Othercontaminant
A- Balla st water; B-Hu ll fouling; C-Ot herstowaway
A-Canals and artificial waterways; B-Tunnelsand bridge
s
C

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counted as TMS as they had not been found in the
natural environment.
Results
Seventeen transplanted mollusc species (TMS) were
recognized in South America (Online Resources 4,
5): ten bivalves (only one freshwater species) belong-
ing to six families (Fig.2) and seven gastropods (four
freshwater and three terrestrials) belonging to three
families (Fig.3).
Transplanted mollusc species were recorded in
16% (31/189) of the ecoregions, specifically in 36%
(10/28) of the marine ecoregions, 23% (12/52) of
the freshwater ones, and 8% (9/109) of the terrestrial
ones. The ecoregions with TMS (31) are generally
located in the peripheral zones of the continent, typi-
cally adjacent to other ecoregions with TMS (Figs.2,
3).
The marine ecoregions with most transplanted
species (Table 1) are Southern Caribbean (3 TMS)
and Uruguay-Buenos Aires Shelf, Northeastern Bra-
zil and Southeastern Brazil (2 TMS each). In fresh-
water, Central Andean Pacific Slopes ecoregion has
three TMS and Western Amazon Piedmont two TMS,
and in the terrestrial environment the only region
with more than one TMS is Alto Paraná Atlantic
Forest with two. In several localities, certain species
were recognized in captivity (Figs.2, 3). The coun-
try with the most TMS is Chile, with five in the wild
and three in captivity, followed by Argentina, Brazil
and Colombia with four, Ecuador with three, Peru
with two, and Uruguay and Venezuela with one each
(Online Resources 4 and 5).
The freshwater Pomacea canaliculata (Lamarck,
1822) is the transplanted gastropod recorded in the
largest number of ecoregions (seven). For bivalves, it
is the marine scissor date mussel Leiosolenus arista-
tus (Dillwyn, 1817), present in four ecoregions. The
transplantation mechanism (Table2) was Movement
of Commodity for all gastropod species except P.
canaliculata, while for bivalves this mechanism cor-
responds to only four of the cases. The remaining
bivalve transplants (six) occurred by Vector, as sev-
eral species are in the two subcategories ‘hull fouling’
and ‘ballast water’.
In South America, TMS are known to cause some
type of impact, both on the natural environment
(e.g., modification of the environment, displacement
of native fauna), and socio-economic (e.g., agricul-
ture, aquaculture, parasite vector) (Table 3, Online
Resouces 4 and 5).
Nine species have undergone modifications in their
historical distributions, but do not fit the transplanta-
tion definition (Table4, Online Resource 6). Further-
more, species such as Mytilus platensis d’Orbigny,
1842 and Perna perna (Linnaeus, 1758) are still
undergoing debate regarding their taxonomy and their
status as natives (Online Resource 6).
Discussion
Given our current knowledge of climate change, it is
expected that more changes in species distribution
and abundance within South America will take place
in the coming decades (McDowell etal. 2014; Bár-
cena etal. 2020). Those changes will be interacting
with other stressors, such as habitat loss, change in
land use, eutrophication, and impacts of invasive spe-
cies. To understand the overall impacts of these fac-
tors, their interactions must be examined, and prob-
ably the first step is to precisely identify the current
distributions of species in the continent.
Species are naturally dispersed by different mecha-
nisms, but human activities have become a common
means of transport, not only moving species from one
geographic region to another, but also expanding their
natural distributions. Studies of non-native species
are many, particularly biased towards invasive species
with noticeable impact on human welfare. However,
the transplantation (as here defined) of species is not
frequently considered, despite their potential socio-
ecological impacts.
Knowledge of the native and non-native mollusc
fauna of South America is heterogeneous. Studies
on mollusc diversity are incomplete in vast regions
of South America (Darrigran etal. 2020) and there-
fore the number of valid native species is not yet well
established (e.g., Salvador 2019; Dos Santos et al.
2021). This lack of knowledge is evidenced in the
present study by the number of transplanted mollusc
species (TMS) recognized for all of South America.
Although there is no up to date record of mollusc spe-
cies present on the continent, the reported mollusc
richness of the largest of the 12 countries in South
America, Brazil, is close to 2,950 species considering

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Chile: Puerto Montt and Chiloé,
X Region (von Brand et al. 2016)
Argopecten purpuratus (Lamarck, 1819)
Argentina: Bahía Blanca,
Buenos Aires (Fiori et al. 2012)
Brazil: Cearà, Baía, Río de Janeiro,
Sao Paulo, Paraná and Santa Catarina
(Teixeira and Creed 2020); Espiritu
Santo (Gomes et al. 2014)
Ecuador: Champion Island, Galápagos
(Carlton et al. 2019)
Leiosolenus aristatus (Dillwyn, 1817)
Colombia: Cartagena Bay,
colombian Caribbean coast (Puyana
et al. 2012; De La Hoz Aristizábal, 2013)
Venezuela: Unare, Caribbean coast
(Lodeiros et al. 2021)
Mytilopsis cf. sallei (Récluz, 1849)
Brazil: Recife (Fernandes et al. 2018)
Mytilopsis trautwineana (Tryon, 1866)
Colombia: Caribe, rivers that dranage
to Caribean coast (Aldridge et al.
2008)
Mytilus platensis d’Orbigny, 1842 Perna perna (Linnaeus, 1758)
Tawera elliptica (Lamarck, 1818)
Chile: Reñaca (Oliva and Durán 2012)
Barnea truncata (Say, 1822)
Mytella strigata (Hanley, 1843)
Brazil: Bombinhas, Palhoça and
Florianópolis, Santa Catarina
(dos Santos et al. 2019)
Colombia: Cartagena, Bolívar (Gracia
et al. 2011)
Chile: Punta Arenas (GBIF 2022)
Uruguay: Cerro Verde (Carranza and
Borthagaray 2009)
Anodontites trapesialis (Lamarck, 1819)
Brazil: artificial lakes, Paraná,
São Paulo, Santa Catarina and
Amazonas (Silva-Souza and Guardia
Felipi 2014)
Peru: coast of Piura and Lambayeque
(Ramirez et al. 2003)

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the freshwater, terrestrial, and marine environments
(Amaral and Jablonski 2005; Simone 2006; Salva-
dor 2019); 1800 species have been recorded in Chile
(Aldea et al. 2019), while approximately 1300 spe-
cies are reported from Argentina (Rumi etal. 2008;
Bigatti and Signorelli 2018; Dos Santos etal. 2021)
and the richness in tropical countries is expected
to greatly exceed those numbers. This information
allows us to affirm that the 17 TMS identified here
for all South America constitute a very conservative
estimate. It is likely that our partial knowledge of the
distribution and taxonomy of many of the molluscs
is masking other potential TMS. Furthermore, there
are also species that are mobilized by humans for
commercial use, cultivation, or research reasons, but
are only found in captivity and have not established
themselves in the natural environment (Darrigran
et al. 2020); those species can potentially become
new transplants in the future.
Darrigran et al. (2020) recognized four zones in
South America based on the highest number of non-
native mollusc species recorded after 1970 and the
associated anthropogenic factors (e.g., urbanization
and trade). These zones are predominantly periph-
eral mainland areas (i.e., Subtropical Atlantic, North-
ern Andes, Central Andes, and Southern Andes) and
were recognized as non-native mollusc hot-spots and
entry points of non-native mollusc species (Darrigran
etal. 2020). Our results show that of the 31 ecore-
gions with TMS, 14 are included in these four hot-
spots of non-native mollusc species. In addition, all
terrestrial ecoregions in which transplanted terrestrial
molluscs were recognized belong to the category of
highest conservation priority at a regional scale (Din-
erstein etal. 1995).
Consequently, it can be deduced that the envi-
ronmental and socioeconomic characteristics that
facilitate the introduction and settlement of non-
native species of molluscs are also correlated with
the transplantation of species within South America
(Table1). This result points to a relationship between
the features of the environment (invasibility) and
those of the transplanted mollusc species (invasive-
ness) (Hicks 2004). Strayer (2022) remarked that the
impacts of transplanted species can be predicted from
(a) the species traits, (b) how axial the translocation
is (i.e., if an existing population is increased, or re-
established at a recently occupied site, or introduced
to a new site), and (c) the rigor of translocation proto-
cols in cases where the aim is conservation or biocon-
trol (Novak etal. 2021).
Two highly successful mollusc species trans-
planted within South America (Pomacea canalicu-
lata and Leiosolenus aristatus) exhibit behavioral and
physiological characteristics that have facilitated their
introduction in other continents. Leiosolenus arista-
tus is a boring species from the western Atlantic and
the Pacific, which was detected off the coasts of Rio
de Janeiro and São Paulo, southeast Brazil (Simone
and Gonçalves 2006). It has also been introduced
to the Cape Verde Islands (Lopes 2011) and to the
Mediterranean coast of Catalonia (López Soriano and
Quiñonero Salgado 2019). Pomacea canaliculata is
among 100 of the worst invasive species worldwide
(Lowe etal. 2000). This species causes major eco-
nomic and public health problems in much of South-
east Asia (Joshi et al. 2017), Europe (López et al.
2010), and Africa (Buddie etal. 2021). Trade, mari-
time routes, and tourism have increased in the twen-
tieth and twenty-first centuries (IUCN 2009; Hobbs
et al. 2018), and are the most important factors in
the introduction of non-native species globally. This
is more pronounced for maritime transport (IUCN
2009) and, accordingly, the majority of TMS in South
America have been recorded from marine ecoregions.
Among the terrestrial molluscs, Darrigran et al.
(2020) considered the slug Sarasinula plebeia (Fis-
cher, 1868) a cryptogenic species. However, Gomes
and Thomé (2004) considered there to be a consensus
that it was introduced into Australia and the several
islands on the Pacific by commerce in the Indian and
Pacific Oceans and that it is probably originally from
tropical areas of America, as also stated by Cowie
etal. (2008). Daglio etal. (2020) noted that Vaginula
behni Semper, 1885, which is currently a synonym of
S. plebeia according to Thomé (1993), was originally
described based on material from Rio de Janeiro, Bra-
zil, indicating that it has been present in South Amer-
ica for a long time, despite being originally described
based on material from New Caledonia, in the Pacific
region. According to Daglio etal. (2020), S. plebeia
Fig. 2 Distribution of transplanted bivalves in South America
by ecoregions (limits of ecoregions in light blue). Ecoregions
that include the known historical geographic distribution are
shown in green; those that include transplant locations are
shown in blue. For each species the location and reference of
transplants are mentioned. More information is available in
Online Resource 4
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Chile: in captivity (PNUD 2014)
Aplysia dactylomela Rang, 1828 Cerithium litteratum (Born, 1778)
Chile: in captivity (PNUD 2014)
Chile: in captivity (Baez et al. 2014)
Cyphoma gibbosum (Linnaeus, 1758) * Pomacea bridgesii (Reeve, 1856)
Argentina: in captivity (Valdez and Cruz
López 2014)
Brazil: Guapimiim (Thiengo personal
communication), hydroelectric power
station Paracambí (Fernandes personal
communication), RJ; Juiz de Fora, MG.
Chile: in captivity Santiago (Letelier et
al. 2007; PNUD 2014)
Perú: along the Peruvian coast, in
aquariums and semi-controlled
environments (Ramirez et al. 2003)
Venezuela: in captivity (Ojasti 2001)
Pomacea canaliculata (Lamarck, 1822)
Argentina: Desaguadero, Colorado, south
Bonaerensean drainages and arheic
waterbodies (Seuffert and Martín 2020)
Chile: Laguna Conchalí, Los Vilos (Letelier
and Soto-Acuña 2008; Jackson and
Jackson 2009; Letelier et al. 2016;
Almendras García 2021)
Ecuador: wide distribution (Correoso et al.
2015, 2017)
Perú: in coastal sites (Ramirez et al. 2003)
Venezuela: in captivity (Ojasti 2001)
Pomacea diffusa Blume, 1957
Argentina: in captivity (Martin 2017)
Ecuador: near Huapuno River, Pastaza
Uruguay: in captivity (Röhrdanz
Rosa 2017)
Venezuela: free in disturbed environmets
(Agudo-Padrón personal communication)
Pomacea maculata Perry, 1810
Peru: coast region, center and south
(Ramírez et al. 2003)
Colosius confusus Gomes, Robinson,
Zimmerman, Obregón & Barr, 2013
Colombia: Neira, Caldas
Ecuador: Guayabilla, Ibarra, Imbabura,
Volcán Pululahua, Pichincha, Papallacta,
Imbabura, Caranqui, Cotopaxi, Otonga,
Chimborazo, Alausí, Tungurahua, Patate,
Bolívar; Napo, Cuyuja, Cosanga, La Florida
(Gomes et al. 2013)
Bulimulus bonariensis (Rafinisque, 1833)
Argentina: Misiones, Salta, Chaco,
Santiago del Estero, Corrientes,
Córdoba, Tucumán, Salta, Jujuy
Brazil
Uruguay: Montevideo Department
(Scarabino 2003)
Sarasinula plebeia (P. Fischer, 1868)
Argentina: Selvas del Río de Oro, Chaco;
Puerto Iguazú, Misiones (Daglio et al.
2020)

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has been reported in Brazil (since 1885), Colombia
(1978), Venezuela (1992), Chile (1993), Ecuador
(2008), Peru (2015), and Argentina (2020); they con-
sidered its presence in Argentina as a transplanted
species.
It is important to note, however, that the geograph-
ical origin of some species remains uncertain. Such
is the case, as mentioned before, for the brown mus-
sel Perna perna, which has been considered native to
either South America (Darrigran etal. 2020) or the
western coasts of Africa (Oliveira etal. 2017). Cala-
zans etal. (2021) clarified the status of P. perna by
analyzing the genetic structure of several populations,
showing that it is native to both continents: popula-
tions in Brazil are derived from natural geographical
expansion from the populations in northern Africa,
which took place over 2000years ago. In addition,
taxonomic problems often prevent proper assess-
ment of the distribution of species. For example, the
geographic range of Mytilus platensis is not reliably
defined, given its complex identification and diagnos-
ability from other Mytilidae, such as Mytilus chilensis
Hupé, 1854 from the southeastern Pacific (Zbawicka
etal. 2018). Some studies indicate that the distribu-
tion of M. platensis extends from southern Brazil to
southern Argentina (Dellatorre etal. 2007), with the
North Patagonian gulfs probably being the southern
limit of its distribution. A characteristic of this spe-
cies is its ecology in the coasts of northern Argentina
and southern Uruguay, where in addition to being
part of the intertidal and infralittoral community, they
form deep banks away from the coast, between 40 and
55m deep (Penchaszadeh 1980). Adding to the com-
plexity is the introduced Mediterranean Mytilus gal-
loprovincialis Lamarck, 1819 in the Puerto Madryn
area (42° 51′ S, 65° 15′ W), where it has hybridized
with the local M. platensis (Zbawicka etal. 2018).
All these caveats being noted, we found that TMS
occur in 18% of the ecoregions in South America,
31% of which are contiguous. Contiguous ecoregions
may have similar or at least favorable environmen-
tal characteristics for the presence of TMS. Most of
these scenarios correspond to contiguous freshwa-
ter ecoregions in line with the hypothesis of Bianco
(1995), that river basins are the main geographic axes
of introduction of non-marine aquatic species. In gen-
eral, TMS transportation routes in South America are
related to economic purposes, namely aquaculture,
agriculture, horticulture, and trade of ornamental
species. Sindermann (1993) warned about the trans-
plantation and introduction of marine species world-
wide in response to the perceived need of expanding
aquaculture industries (e.g., fish, shrimp, bivalve mol-
luscs). On the other hand, Bartley etal. (2005) stated
that the use of non-native species is a proven means of
increasing productivity and value of aquatic ecosys-
tems. Likewise, FAO (2018) promoted practices for
the introduction of new species and the responsible
movement of live aquatic animals. According to the
aforementioned, Álvarez León (2014) pointed out that
it is necessary to identify the impacts of transplanting
native species and introducing non-native species, in
addition to regulating their use in each country. For
instance, most of the transplants in Colombia have
originated from aquaculture (Restrepo-Santamaría
and Álvarez-León 2013) and, as the technological
packages of some species were developed, they began
to be used in regions different from their basins of ori-
gin (“trans-introductions” according to Bianco 1995),
highlighting the importance of this type of pathway
for the transplantation of species. For example, Ano-
dontites trapesialis (Lamarck, 1819) was transplanted
to the north coast of Peru for aquaculture purposes
in integrated fish and duck cultures (Ramírez et al.
2003). Agudo-Padrón (2019, 2022) reported this spe-
cies as a contaminant in fish farming tanks, farming
areas, and tourist fishing facilities in rivers in south-
ern Brazil. The life cycle of A. trapesialis includes
an ectoparasitic larval stage that adheres to the gills
and fins of host fish. The cycle is completed in the
fish rearing tanks, which has a negative impact on
the development of this resource. Tanks are mainly
contaminated through the inadvertent introduction
of infested fish. Likewise, Agudo-Padrón (2005)
reported that the Federal Rural University of Pernam-
buco experimented with freshwater fish farming with
Fig. 3 Distribution of transplanted gastropods in South Amer-
ica by ecoregions (limits of ecoregions in light blue). Ecore-
gions that include the known historical geographic distribution
are shown in green; those that include transplant locations are
shown in blue. The blue stars indicate specimens in captivity
(not considered TMS). For each species the location and refer-
ence of transplants are mentioned. *The distribution of Poma-
cea bidgesii is difficult to establish because Pomacea diffusa
was previously considered a subspecies of P. bridgesii and,
although they are currently recognized as separate species,
misidentification of P. diffusa as P. bridgesii is frequent (Cowie
and Thiengo 2003; Hayes etal. 2008, 2009; Cowie etal. 2017).
More information is available in Online Resource 5
◂

G.Darrigran et al.
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A. trapesialis for human consumption, using host fish
such as carp, Cyprinus carpio (Linnaeus, 1758) and
tilapia (Oreochromis sp.).
Although no further records of freshwater bivalve
culturing were detected in South America, there have
been instances of culturing the gastropods Pomacea
bridgesii (Reeve, 1856) and Pomacea lineata (Spix
Table 1 Number of transplanted mollusc species (TMS) according marine, freshwater and terrestrial ecoregions of South America
Following Darrigran etal. (2020): NNMS, number of non-native mollusc species (from outside South America); Hotspots of NNMS
according to Darrigran etal. (2020)
*Ecoregions outside the hotspots of non-native mollusc species; nc, ecoregion not considered in Darrigran etal. (2020)
Ecoregion TMS NNMS Hotspots NNMS
Marine environment
Southern Caribbean 3 1 Northern Andes
Northeastern Brazil 2 3 *
Uruguay-Buenos Aires Shelf 2 5 Subtropical Atlantic
Southeastern Brazil 2 6 Subtropical Atlantic
Central Chile 1 4 Southern Andes
Chiloense 1 4 Southern Andes
Channels and Fjords of Southern Chile 1 1 *
Eastern Brazil 1 2 *
Eastern Galapagos Islands 1 nc nc
Southwestern Caribbean 1 1 Northern Andes
Total marine ecoregions with TMS 10/28 = 36%
Freshwater environment
Central Andean Pacific Slopes 3 7 Central Andes
Western Amazon Piedmont 2 nc *
South Andean Pacific Slopes 1 3 Southern Andes
North Andean Pacific Slopes Rio Atrato 1 3 *
Mar Chiquita – Salinas Grandes 1 4 *
Amazonas High Andes 1 2 *
Paraíba do Sul 1 5 Subtropical Atlantic
Fluminense 1 6 Subtropical Atlantic
Cuyan—Desaguadero 1 2 *
Upper Paraná 1 6 Subtropical Atlantic
Orinoco Llanos 1 3 Northern Andes
Orinoco Guiana Shield 1 1 *
Total freshwater ecoregions with TMS 12/52 = 23%
Terrestrial environment
Alto Paraná Atlantic forests 2 16 Subtropical Atlantic
Southern Andean Yungas 1 16 *
Magdalena Valley montane forests 1 13 Northern Andes
Napo moist forests 1 6 *
Eastern Cordillera real montane forests 1 5 *
Northwestern Andean montane forests 1 3 *
Dry Chaco 1 12 *
Humid Chaco 1 7 *
Uruguayan savanna 1 22 Subtropical Atlantic
Total terrestrial ecoregions with TMS 9/109 = 8%

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in Wagner, 1827). Souza etal. (2013) mentioned that,
because of their high rates of reproduction, growth,
and early sexual maturity, as well as for having been
previously used as food by settlers in Trinidad, Guy-
ana, French Guiana, and northern Brazil, they were
cultivated to evaluate the viability of their use as a
Table 2 Pathways of transplanted mollusc species in South America (seven Gastropoda and ten Bivalvia species)
The mechanisms and main categories are based on Faulkner etal. (2020); *, gastropod species; **, bivalve species. More than one
pathway sub-category wasrecognized for some species
#The identification of Pomacea bridgesii is suspect, probably mistaken for Pomacea diffusa; P. diffusa is widely distributed through
much of the Amazon basin and is found around the world in the aquarium trade, while P. bridgesii is restricted to Bolivia and the
western Amazon basin (Hayes etal. 2008, 2009; Cowie etal. 2017)
Mechanism of Transplant Pathway Category Pathway Sub-category
Movement of commodity Escape Aquaculture
1. Anodontites trapesialis** Ramírez etal. (2003)
2. Argopecten purpuratus** von Brand etal. (2016)
3. Mytilopsis trautwineana** Albridge etal. (2008)
4. Pomacea maculata* Alcantara-Bocanegra and
Nakagawa-Valverde (1996)
5. Tawera elliptica** Oliva and Durán (2012)
Ornamental
1. Pomacea diffusa* Scarabino etal. (2012)
Martín (2017)
Horticulture and Agriculture
1. Bulimulus bonariensis* Frana and Massoni (2011)
2. Colosius confusus* Gomes etal. (2013)
Live bait
1. Pomacea canaliculata* Seuffert and Martín (2020)
Live food
1. Pomacea canaliculata* Seuffert and Martín (2020)
Other escape
1. Pomacea bridgesii*#Martins and Alves (2008)
2. Pomacea canaliculata* Seuffert and Martín (2020)
3. Sarasinula plebeia* Daglio etal. (2020)
Contaminant Parasite of animals
1. Anodontites trapesialis** dos Santos Silva etal. (2021)
Vector Stowaway Hull fouling
2. Barnea truncata** Fiori etal. (2012)
3. Leiosolenus aristatus** Carlton etal. (2019)
4. Mytella strigata** Lodeiros etal. (2021)
5. Mytilopsis cf. sallei** Queiroz etal. (2020)
6. Mytilus platensis** Cárdenas etal. (2020)
7. Perna perna** PNUD (2014)
8. Pomacea canaliculata* Seuffert and Martín (2020)
Ballast water
1. Barnea truncata** Fiori etal. (2012)
2. Mytella strigata** Lodeiros etal. (2021)
3. Mytilopsis cf. sallei** Queiroz etal. (2020)
4. Mytilus platensis** Cárdenas etal. (2020)
5. Perna perna** PNUD (2014)

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possible new food source. However, concerning the
identification and geographical range of both species
we highlight that P. bridgesii is restricted to Bolivia
and the western Amazon basin, whereas Pomacea dif-
fusa Blume, 1957 is widely distributed through much
of the Amazon basin and it is the one in the aquarium
trade that is found around the world now (Cowie etal.
2017). Regarding P. lineata, it occurs primarily in the
Atlantic drainages of Brazil—the Brazilian Atlantic
Forest biogeographic province—and is replaced by
its close relative Pomacea dolioides (Reeve, 1856) in
northern South America (Surinam, French Guiana,
Guyana, and Venezuela). Thus, the reference to P.
lineata is probably a misidentification of P. dolioides
(Thiengo etal. 2011; Cowie etal. 2017).
In short, the growing circulation of people, the
increase in trade, and changes both to the environ-
ment and in production practices, give rise to new
threats to production systems and to the natural
environment (FAO 2015). In general, non-native spe-
cies can have negative impacts on recipient environ-
ments and so do transplanted species (third premise
of Material and Methods). According to Simberloff
et al. (2012), when non-native species invade, they
can modify community structure and ecosystem func-
tion, becoming ecosystem engineers (Jones et al.
1994), increasing the risk of extinction for native
species (Collado 2016; Collado etal. 2019), causing
negative effects on human health, and becoming a
serious socio-economic threat (Pejchar and Mooney
2009). For example, Colosius confusus Gomes etal.
2013 occurs in the Andean region, in regions for
production of cut flowers and coffee in Colombia
and Ecuador (Gomes etal. 2013). In northern Peru
it was found attacking coffee in Canchaque. It is
also found in two other Departments in Peru (Ama-
zonas and Cajamarca), where it is sympatric with
native species in urban areas. Although not causing
Table 3 List of mollusc species transplanted in South America and their environmental impacts and positve (+) or negative (−)
socioeconomic effects
A (−), losses in agriculture; AQ (+), aquarium trade; D (−), species displacement; E (−), environment modification; F (−), losses in
aquaculture; P (+), profits from aquaculture; V (−), parasite vector. Details in Online Resource 4 and 5
#The identification of Pomacea bridgesii is suspect, probably mistaken for Pomacea diffusa (Hayes etal. 2008, 2009; Cowie etal.
2017)
Impact on natural environment Socioeconomic impact
Meaningful Moderate Null or
unknown
Meaningful Moderate Null or
unknown
Gastropoda
Bulimulus bonariensis (Rafinisque, 1833) x A
Colosius confusus Gomes etal., 2013 x A
#Pomacea bridgesii (Reeve, 1856) D AQ
Pomacea canaliculata (Lamarck, 1822) D–E A–V
Pomacea diffusa Blume, 1957 X AQ
Pomacea maculata Perry, 1810 D–E x
Sarasinula plebeia (Fischer, 1868) x A-V
Bivalvia
Anodontites trapesialis (Lamarck, 1819) x F
Argopecten purpuratus (Lamarck, 1819) x P
Barnea truncata (Say, 1822) E x
Leiosolenus aristatus (Dillwyn, 1817) D x
Mytella strigata (Hanley, 1843) D F
Mytilopsis cf. sallei (Recluz, 1849) x x
Mytilopsis trautwineana (Tryon, 1866) x F
Mytilus platensis d’Orbigny, 1842 D x
Perna perna (Linnaeus, 1758) x P
Tawera elliptica (Lamarck, 1818) x P

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agricultural damage as in Colombia and Ecuador,
and exhibiting a different external coloration in rela-
tion to the forms found attacking coffee, no genetic
variation was found between them. Colosius confusus
was first collected in Peru in 1970 and in Ecuador in
1985 (Gomes etal. 2013). In Ecuador, it is the second
most commonly found Andean species of Veronicelli-
dae after Colosius pulcher (Colosi, 1921). Also, even
after Colosi (1921, 1922) examined many specimens
from Ecuador, he did not describe C. confusus, which
lends support to it being a recent transplant. The lack
of variation observed in C. confusus specimens from
Ecuador, Colombia, and Peru using mitochondrial
DNA sequences is consistent with recent transplanta-
tion between countries. Gomes etal. (2013) indicated
thus that the species is probably originally from Peru.
In Ecuador, the production of cut flowers began in the
late 1970s with exports in the early 1980s (Acción
Ecológica 2000), indicating that the spread of C.
confusus could be related to expanding commercial
flower production.
The terrestrial snail Bulimulus bonarien-
sis (Rafinesque, 1833) was originally described
based on material from Buenos Aires but spread to
northeastern Argentina (Cuezzo etal. 2013), where
it is reported as a pest of soybeans (Frana and Mas-
soni 2011). It is also reported from Uruguay (Scara-
bino 2003) and Brazil, to both of which it was prob-
ably transplanted considering its pest status. The
case of Bulimulus species is remarkable and a good
example of how necessary it is to expand taxonomic
work based not only on molecular studies but also
on morphology. Maintaining taxonomic collections
in public institutions is important for understanding
historical distribution changes as well as the spread
of certain species because of the changes produced
by people. Bulimulus apodemetes (d’Orbigny, 1835)
has an unusually wide distribution for a terrestrial
gastropod and the marked intraspecific variabil-
ity of its shells suggests a complex of species. The
advance of agricultural frontiers in some areas of
the Yungas (Bolivia, Peru and Argentina) and Dry
Chaco (Bolivia, Paraguay and Argentina) ecore-
gions has produced the appearance of specimens
similar to B. apodemetes in areas where they did not
occur before. However, these abundant specimens
in soybean crops and other grains have not yet been
correctly identified. New studies are needed that
Table 4 Mollusc species with alterations in their known historical distribution, but which are not considered transplantations
The causes of the alterations were classified as: 1, Nomenclatural and/or taxonomic issues; 2, natural dispersal, without direct human
intervention; 3, species intercepted in trade traffic; 4, species mobilized by humans for repopulation in their known historical geo-
graphic distribution. Details of the distribution of these species in Online Resource 6
*Dispersal due to El Niño Southern Oscillation
Species References Cause of
altera-
tions
GASTROPODA
Terrestrial Bulimulus apodemetes (d’Orbigny, 1835) Miquel (1991), Dos Santos etal. (2021) 1
Gastrocopta servilis (Gould, 1852) Doering (1874), Pilsbry (1916–1918), Parodiz (1957), Fernández
(1973)
1
Orthalicus pulchellus (Spix, 1827) Cruz (1995), Simone (1999) 1
Aplysia juliana Quoy & Gaimard, 1832 Soto (1985), Tomicic (1985), Castilla etal. (2005), Uribe etal.
(2013)
2*
Marine Monoplex keenae (Beu, 1970) Beu and Cernohorsky (1986), Ashton etal. (2008), Araya (2015) 2*
Monoplex wiegmanni (Anton, 1838) Castilla etal. (2005), Ashton etal. (2008) 2*
BIVALVIA
Marine Modiolus carvalhoi Klappenbach, 1966 Zaffaroni (2000); Scarabino etal. (2006), Silveira etal. (2006),
Borthagaray and Carranza (2007), Huber (2010)
2
Semimytilus patagonicus (Hanley, 1843)
[= S. algosus (Gould, 1850)]
Alamo and Valdivieso (1987, 1997), Coan and Valentich-Scott
(2012), Bigatti etal. (2014), Signorelli and Pastorino (2021a, b)
3
Freshwater Diplodon chilensis (Gray 1928) Parada and Peredo (2005), Peredo etal. (2005), Valdovinos and
Pedreros (2007), Fuentealba etal. (2010)
4

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focus on this malacofauna associated with human
environments.
The movement of species is among the most seri-
ous environmental threats of the new millennium, as
the relocation of species beyond their native or his-
torical geographical range has intensified in the last
five decades; however, its general effects on the envi-
ronment are not yet sufficiently understood (Anton
etal. 2019). Since the impacts of TMS may be high,
both for natural environments and for socioeconomic
systems, we emphasize the need for further studies of
transplantations in South America and elsewhere.
Acknowledgements The authors are grateful for the sugges-
tions and help provided by Robert Cowie and the comments of
the two anonymous reviewers. GD, DEGG, and CD were par-
tially supported by Project PICT2019-01417, Agencia Nacional
de Promoción Científica; GD, CD by Universidad Nacional de
La Plata (11/N927), GD, PEP and DEGG (PIP1966) and MGC
(PIP0050) by Consejo Nacional de Investigaciones Cientifícas
y Técnicas
Funding Agencia Nacional de Promoción Científica
(PICT2019-01417); Universidad Nacional de La Plata (11/
N927); Consejo Nacional de Investigaciones Científicas y Téc-
nicas (PIP1966; PIP0050).
Availability of data and materials All data analyzed during
this study are included in this published article and its supple-
mentary information files.
Code availability Not applicable.
Declarations
Conflict of interest The authors have no conflicts of interest
to declare.
Ethics approval Not applicable.
Consent to participate All authors declare consent.
Consent for publication All authors declare consent.
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