Biodivers Conserv (2010) 19:3471–3487

DOI 10.1007/s10531-010-9908-8

ORIGINAL PAPER

Temporal trends in non-indigenous freshwater species

records during the 20th century: a case study
in the Iberian Peninsula

Fernando Cobo • Rufino Vieira-Lanero • Enrique Rego •

Marı́a J. Servia

Received: 16 March 2010 / Accepted: 6 August 2010 / Published online: 24 August 2010
Ó Springer Science+Business Media B.V. 2010

Abstract Galicia (NW Spain) is a region with a high number of freshwater endemics,
and probably the best preserved area concerning fish populations in the Iberian Peninsula,
where records of non-indigenous freshwater species are recent when compared to the rest
of the Peninsula. Detailed analysis of introductions of those species with records after 1900
present in both areas shows that delays were up to 100 years for species introduced on the
Iberian Peninsula at the beginning of the twentieth century, but the tendency adjust to a
decreasing linear regression, with species introduced after 1995 being almost immediately
present in Galicia. We underline the outstanding role of aquarium trade on these results.
Analysis of temporal trends highlights several periods with numerous introductions, and
shows a different trend in the last decade depending on the group of organisms, with a clear
deceleration in introduction rates of vertebrates, but a continuous growing trend for
invertebrates. Recent educational programs might be responsible for the reduction in the
inflow of vertebrates, but there is still a need for the control of less conspicuous but equally
harmful invertebrates and plants, as it will take longer to make both stakeholders and
public aware of their detrimental effects on their new habitats.

Keywords Non-indigenous species  Freshwater habitats  Temporal trends  Galicia 

Iberian Peninsula  Aquarium trade

F. Cobo  R. Vieira-Lanero
Departamento de Zooloxı́a e Antropoloxı́a Fı́sica, Universidade de Santiago de Compostela,
Campus Sur s/n, 15782 Santiago de Compostela, Spain

F. Cobo  R. Vieira-Lanero
Estación de Hidrobioloxı́a ‘‘Encoro do Con’’, Castroagudı́n s/n, 36617 Vilagarcı́a de Arousa,
Pontevedra, Spain

E. Rego
Avda. das Carolinas 50, 36600 Vilagarcı́a de Arousa, Pontevedra, Spain

M. J. Servia (&)
Departamento de Bioloxı́a Animal, Bioloxı́a Vexetal e Ecoloxı́a, Facultade de Ciencias,
Universidade da Coruña, Campus da Zapateira s/n, 15008 A Coruña, Spain
e-mail: mservia@udc.es

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Introduction

Organisms have been moved around the world as consequence of human activities
throughout History, but some of these species have severely affected their new habitats,
changing the composition of the original communities and even their physical habitat.
Thus, one of the many examples of the initiatives promoted to achieve a better knowledge
on alien species is the European program SEBI2010 (Streamlining European 2010 Bio-
diversity Indicators). This program includes invasive alien species among the set of bio-
diversity indicators used to assess and inform about the status of biodiversity in Europe, as
they constitute an important threat (EEA 2007).
Freshwater ecosystems have been the recipients of many alien species, both by unin-
tentional or deliberate introductions (Gherardi et al. 2008). Some areas, such as the North
American Great Lakes or central Europe, have been reported to be severely affected by
freshwater invasions, mainly due to the existence of inland navigational routes (Ojaveer
et al. 2002). Contrastingly, despite the decades invested in taxonomic and faunistic studies,
experts state that it is still difficult to get a good overview of biodiversity of freshwater
ecosystems, as available knowledge is largely scattered and focused on a few well-studied
groups (Balian et al. 2008). As a consequence, the presence of Non-Indigenous Freshwater
Species (NIFS) might introduce important biases on initiatives promoting the biodiversity
assessment of certain areas, mainly in groups where inadvertent introductions are frequent
(Strong et al. 2008), as they might be considered as indigenous species, or they might be
responsible of the extinction of many others.
NIFS have already arrived to areas where potential for introduction is apparently low.
Galicia (NW Spain) (Fig. 1) is a region that presents clear natural boundaries that isolate it
from the rest of the Iberian Peninsula (IP), with the river Miño on the South and a mountain
range on the East. It has a dense river network, with numerous small coastal rivers with
high slope and, except for the lower part of the river Miño, none of them are navigable.
Besides, dams are more recent and smaller than those constructed in the rest of Spain, with
a delay of almost 15 years on the average date of construction (estimation done using the
database of the Spanish Dams and Reservoirs Society, SEPREM 2010). All these reasons
might help to explain why detection of many NIFS in Galicia is recent when compared to
the rest of the IP, as authors have established a clear association between navigation and
river impoundment with the presence of NIFS (see for example Clavero et al. 2004; Filipe
et al. 2004; Durán and Anadón 2008; Ribeiro et al. 2008).
Although the presence of NIFS in Galicia is recent, researchers have underlined their
steep increase on recent years, and alert against their potential detrimental effects (Gon-
zález and Cobo 2006). Concerns arise as this region presents both a high biodiversity and a
high number of endemic species linked to freshwater habitats. Thus for example, although
estimates on the number of endemic invertebrates are difficult to make due to the lack of
complete inventories (Sánchez-Fernández et al. 2008), medium altitude streams of the
eastern part of Galicia have been highlighted as habitats with a very rich fauna of aquatic
Coleoptera, in particular endemic species, and they have been considered among the
habitats to have the highest conservation value in what refers to Iberian aquatic Coleoptera
(Ribera 2000). As a consequence, local population extirpations due to NIFS could repre-
sent global extinctions. Concerning aquatic vertebrates, Galicia is well known for the high
diversity of amphibians (Galán 1999), and it is probably the best preserved area concerning
fish populations in the Iberian Peninsula (Elvira and Almodóvar 2001).
Thus, this paper aims at (i) making a complete compilation of NIFS in a region with
high number of endemics but sparse data on alien species, which represent an obvious

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Fig. 1 Study area, with Galicia (NW Spain) highlighted

threat to those endemics, and (ii) analysing the temporal trends of NIFS records in Galicia
when compared to the IP, as they may help identifying important time intervals and factors
for introductions, which might be useful for the establishment of policies for prevention,
control and mitigation of impacts in similar small watersheds.

Materials and methods

Due to lack of information of NIFS on the area, and with the aim of offering an exhaustive
revision and reducing bias on the results, most of the data used in this study have been
gathered from scientific literature, but we have included also many references of grey
literature and information obtained during extensive sampling campaigns of researchers
working at the Hydrobiological Field Station of the University of Santiago de Compostela
(Galicia, Spain) during recent years (See Appendix 1). For NIFS on the IP, we have used a
recent review by Garcı́a-Berthou et al. (2007), although we have reviewed also grey
literature from Spain and Portugal in order to confirm dates of introduction. As recom-
mended by the European Environment Agency (EEA 2007), organisms included in this
review are grouped into vertebrates, invertebrates (excluding parasites) and macrophytes
(excluding helophytes).

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Appendix 1 lists NIFS detected on natural areas. For year of first record, we maintain
the year of first publication when no other date was found. For temporal analysis, time
intervals such as decades have been replaced with the midpoint. Records that need further
confirmation are highlighted. As supplementary information, translocated species to
Galicia were included when they were introduced from other areas of the IP. We have not
included records already considered as clear failed introductions by the authors (see for
example Ribeiro et al. 2009).
In order to evaluate the importance of introduction vectors a v2 test was performed,
using only those species with known invasive status (categories: invasive, naturalized,
naturalized/invasive. See Appendix 1).
Temporal trends of introductions have been analysed using the cumulative number of
NIFS from 1900 onwards in ten-year intervals; pre-1900 introductions were also estimated
(EEA 2007). Delay of arrival to Galicia with respect to the IP for those species with
records after 1900 is offered as mean ± S.E. We analysed the differences in trends
between both areas using the software TRIM (TRends and Indices for Monitoring data;
Pannekoek and van Strien 2005). TRIM is software for the analysis of time series of counts
that estimates yearly indices and trends, and assess the effects of covariates on them, using
a Poisson general log-linear model (McCullagh and Nelder 1989) and accounting for
overdispersion and serial correlation of the data.
TRIM permits trends to be established in parts of the time series by using change points,
which are time points where a significant change in the slope parameter happens. In order
to get the most parsimoniuous model (a model with as few parameters as possible without
compromising its explanatory power), TRIM implements a stepwise model selection
procedure (Pannekoek and van Strien 2005). This procedure repeats the following steps:
1. Calculation of Wald statistics for the difference of the parameters before and after each
change point and their associated significance levels. If the largest significance level
exceeds a certain threshold value (default probability: 0.20) the corresponding change
point is removed from the model.
2. For all removed change points except the last one, calculation of a score statistic to
assess the significance of the difference in parameters before and after the change
point. If the smallest significance level is smaller than a threshold value (default
probability: 0.15) the change point is added to the model.
The procedure stops if no change points can be either removed or added. We took the
year 1900 as the starting point of the series and included ‘‘Area’’ as a covariate with two
categories (Galicia and IP). We have analysed trends for the whole number of NIFS, as
well as for vertebrates and invertebrates independently. Analysis of trends for macrophytes
was not possible due to the low number of species and their recent introduction.

Results

In terms of numbers, there are clearly less records of non native freshwater species in
Galicia than in the rest of the IP, with a total number of 31 NIFS for Galicia and 88 for the
IP (See Appendix 1). Detailed analysis of introductions of those species with records after
1900 present in both areas shows that medium delay for their arrival to Galicia with respect
to the IP was 12.75 ± 4.64 years for invertebrates, 42.33 ± 27.55 years for vertebrates,
and 56.75 ± 17.38 years for hydrophytes. Two species, Haliplanella lineata and
Xenostrobus securis, were cited for the first time in Galicia (Ramil 1987; Garci et al. 2007).

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120

100

y = -0.9476x + 1890.6
80
R 2 = 0.7659
Delay (years)

60

40

20

-20
1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010
Date of introduction

Fig. 2 Relationship between the date of first introduction in the IP and the delay for introduction in Galicia
(years) of non indigenous freshwater species (NIFS)

A scatter plot (Fig. 2) shows that delays were up to 100 years for species introduced on the
IP at the beginning of the 20th century, but the tendency adjust to a decreasing linear
regression, with species introduced on the IP after 1995 being almost immediately present
in Galicia.
Concerning pathways of introduction in Galicia, statistical analysis shows an out-
standing role of aquarium trade on the arrival of species (v2 = 12.8, d.f. = 4, P = 0.012,
n = 25). Indeed, some of these have become invasive in a short time. Examples are the
slider turtle (Trachemys scripta) (Pleguezuelos 2002), or the macrophytes Egeria densa,
Eichhornia crassipes and Azolla filiculoides, all of them introduced in the last 20 years.
Using the whole number of NIFS in both areas, temporal analysis of their arrival reveals
a steep increase in their number after 1970, both for the total number of species and for
each of the groups (Fig. 3). However, trends of cumulative number calculated using TRIM
software (Fig. 4) show that for total NIFS Galicia presents steeper slopes than the IP during
almost all the studied period (Fig. 4a). Differences are clear for the 1931–1970 periods,
even if Galicia was incorporating less NIFS than the IP (Fig. 3a). However, as trends
reflect changes from the starting point of the series (1900), and the starting number of NIFS
in Galicia was lower than that of the IP, small incorporations are shown as big changes.
Interestingly, however, this tendency persists for the 1981-2009 periods, when number of
NIFS had notably increased.
Independent analyses of trends for invertebrates and vertebrates show that changes in
introduction rates in invertebrates are non significant for Galicia, which means that the
introduction rate has been constant during the whole period (Fig. 4b). As for the IP, the
most significant increase in the introduction rate occurs during 1981–1990. In the case of
vertebrates (Fig. 4c), more significant changes are detected, with important introductions
in the IP during 1901–1910, 1941–1950 and 1991–2000. For Galicia there is a long period
of almost 50 years with no introductions, with the first of the century occurring during
1941–1950. A second important change appears also after a period of 20 years with no
introductions, in 1981–1990. Especially noteworthy are the divergent results obtained for
the two groups during the last decade (2001–2009), since for vertebrates there is a sig-
nificant deceleration on the introduction rates both for the IP and Galicia, while for
invertebrates slopes show no changes in any of the areas.

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a b
90 40
Cumulative number of NIFS

Cumulative number of NIFS

80
IP GALICIA IP GALICIA
70
30
60

50
20
40

30

20 10

10

0 0
1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2009 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2009

c d
20
50

Cumulative number of NIFS

Cumulative number of NIFS

IP GALICIA
40 IP GALICIA

30
10

20

10

0 0
1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2009 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2009

Fig. 3 Cumulative number of non-indigenous freshwater species (NIFS) recorded in the Iberian Peninsula
(IP) and Galicia. (a) Total NIFS, (b) invertebrates, (c) vertebrates and (d) macrophytes

Discussion

Despite the obvious constraint watersheds represent for the spread of freshwater fauna,
introductions of NIFS in the IP have been reported in many papers, mainly concerning
animals (see for example reviews in Elvira and Almodóvar 2001 or Garcı́a-Berthou et al.
2007). In Galicia, at the present moment invertebrate NIFS contribute for 0.91% of the
whole freshwater invertebrate fauna (estimated to amount to over 1320 species; González
and Cobo 2006), while vertebrates associated to freshwater habitats contribute for about
16.7% (total estimated number: 55 species, De Castro 2002) and macrophytes for 5% (total
estimated number: about 120, Romero-Buján 2008). These figures are well under those
already reported for other areas (Gherardi et al. 2008; Keller et al. 2009), and most of these
aliens were not reported to be present in the region until the last two decades. Reasons for
this are diverse. Thus, as mentioned in the Introduction section, most Galician rivers are
not navigable, and rates of river impoundment are well under those of other watersheds. In
addition, we should stress also the fact that, in contrast with other areas of the IP, Galicia
have never suffered from aggressive programs of introductions. This has been the case for
example of lake Banyoles (NE Spain), where numerous species of freshwater fishes have
been introduced since the beginning of the XXth century (Garcı́a-Berthou and Moreno-
Amich 2000). Indeed, nonnative assemblages of northeastern Iberian basins have been
found to be more similar to those of France than to those of the rest of the Iberian
Peninsula, indicating a main introduction route (Clavero and Garcı́a-Berthou 2006).

Fig. 4 Multiplicative slopes (± 95% CIs) standing for the changes in the cumulative number of non- c
indigenous freshwater species (NIFS) recorded in the Iberian Peninsula (IP) and Galicia as calculated by
TRIM (see Methods). (a) Total NIFS, (b) invertebrates and (c) vertebrates

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a 1.8

IP
GALICIA

1.4
Slope

1.0

0.6
1901-10 1911-20 1921-30 1931-40 1941-50 1951-60 1961-70 1971-80 1981-90 1991-00 2001-09

b 2

IP
GALICIA

1.6
Slope

1.2

0.8
1901-10 1911-20 1921-30 1931-40 1941-50 1951-60 1961-70 1971-80 1981-90 1991-00 2001-09

c 3

IP
GALICIA

2.1
Slope

1.2

0.3
1901-10 1911-20 1921-30 1931-40 1941-50 1951-60 1961-70 1971-80 1981-90 1991-00 2001-09

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Certainly, some of the literature we have reviewed may contain inaccuracies, and the
undersampling of some groups, mainly invertebrates and plants, may have changed some
results of our work. Thus for example, an important contribution to the clear increase in
records on the IP during 1981–1990 is the work done on Crustaceans by Paulo and
Moutinho (1983), Forés et al. (1986) and Baltanás et al. (1996). This fact stresses the
sudden changes in records that can occur when important gaps in the knowledge of certain
taxa are filled.
However, at the present moment aquarium trade seems to be the dominant vector of
introduction of NIFS in Galicia, as it has already been shown in other areas (Padilla and
Williams 2004; Rixon et al. 2005; Karatayev et al. 2009; Ribeiro et al. 2009), and it has
been highlighted as a priority target for legislation (Keller et al. 2009; Strayer 2010).
Indeed, the analysis of the delay for the arrival to Galicia of NIFS introduced in the IP after
1900 reveals, as expected, a clear reduction and, interestingly, the decreasing linear
regression that best fits the data intersects the x-axis (delay = 0) around 1995, the same
year the Schengen agreement opened the international borders to the free circulation of
persons and goods. This event has probably facilitated the uncontrolled translocation of
organisms, mainly vertebrates, across international borders, and it could also favour the
arrival of invertebrates and plants, either deliberately or accidentally transported. Actually,
this effect has been observed in Eastern European countries, as their integration to the
European Union has boosted the spread of invasive bird species (Chiron et al. 2010).
However, the Schengen agreement has been linked to social and economic changes, and
more investment on research and therefore more capacity in detecting new species, or more
researcher’s awareness on NIFS problems, might have had an important influence on this
result.
An important outcome of the trends analysis is the deceleration shown for rates of
vertebrate NIFS introductions in the last decade, both for the IP and Galicia. Indeed, our
study shows few new vertebrate NIFS in the list in comparison with those listed by Garcı́a-
Berthou et al. (2007). However, invertebrates show no changes in recent rates of intro-
duction, neither in the IP nor in Galicia, which indicates a continuous growing trend. Both
Portugal and Spain have adopted a number of legislative instruments for the control and
mitigation of impacts of invasive species (See for example in Portugal: Decreto Ley
565/1999, in Spain: Ley 42/2007, del Patrimonio Natural y de la Biodiversidad), which
include the prohibition of trade with exotics known to be invasive. However, there is still
much work to be done for achieving a real control of NIFS trade, as the increasing number
of aquarists and the easy communication among them using the numerous forums on the
web have increased the exchanging of organisms, with the associated risk of escapes or a
final release in natural habitats (Padilla and Williams 2004). Thus, it is probable that the
different tendency observed in invertebrates and vertebrates is due to the influence of
education programs on exotic species, as at least some vertebrates are already perceived by
stakeholders and general public as harmful invasives. Contrastingly, there is still much
work to be done on the less conspicuous but equally harmful invertebrates and plants, as it
will take longer to make both stakeholders and public aware of their detrimental effects on
their new habitats.
A step forward will be to evaluate the invasion risk posed by species used in aquarium
trade, as it has already been done on the Great Lakes (Rixon et al. 2005), as well as to
better understand the impact of NIFS on freshwater endemics. Analysis of trends in
geographical areas where introductions of NIFS are also recent might help to better pin-
point the causes of the problem, as well as to efficiently design protective measures against
new introductions.

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Acknowledgements We thank two anonymous reviewers for the helpful comments on the manuscript.
Facilities needed for this work were provided by the Hydrobiological Field Station ‘‘Encoro do Con’’
(University of Santiago de Compostela). This study was supported by project CGL2009-10868 of the
Ministry of Education and Science (Spain).

Appendix 1

Non-indigenous freshwater species recorded in the Iberian Peninsula and Galicia. Status
and Pathway/Vector only listed for Galicia. NR: No record. **: Records that need to be
confirmed. \: Before. [: After.
Habitat (HB): 1: Streams and rivers (excluding estuaries); 2: Lakes and reservoirs; 3:
Ponds and pools; 4: Rice fields; 5: Estuarine or saline waters.
Status (ST): U: Unknown; N: Naturalized; I: Invasive; N/I: Invasive only in certain
locations.
Pathway/Vector (P/V): A: Angling; AB: Angling, bait; AQ: Aquaculture; AT: Aquar-
ium trade; F: Fouling; FE: Farm escapes; FP: Fouling, secondary dispersal from Portugal;
IR: Intentional release; U: Unknown (Table 1).

Table 1
Group/species First record Reference HB ST P/V

IP Galicia

Cnidaria
Cordylophora caspia 2001 Garcı́a-Berthou et al. (2007) 1,2,5
(Pallas, 1771)
2006 González and Cobo (2006) N F
Craspedacusta sowerbyi 1968 Garcı́a-Berthou et al. (2007) 1,2
Lankester, 1880
1994 González and Cobo (2006) U U
Haliplanella lineata 1987 1987 Ramil (1987) 5 N F
(Verrill, 1869)
Blackfordia virginica 1987 NR Moore (1987) 5
Mayer, 1910
Platyhelmintha
Girardia tigrina (Girard, U Garcı́a-Berthou et al. (2007) 1,2
1850)
2006 González and Cobo (2006) N/I AT
Dendrocoelum lacteum 1993 NR Carranza and Giribet (1997)
(Müller, 1774)
Mollusca
Gyraulus chinensis 1979 NR Garcı́a-Berthou et al. (2007) 2,4
(Dunker, 1848)
Physella acuta 1845 Morelet (1845) 1,2,4
(Draparnaud, 1805)
1932 Pardo (1932) N/I U
(as Physa acuta)
Physella gyrina (Say, 1821) 1988 NR Altaba et al. (1988)
Helisoma duryi seminole 1984 NR Altaba et al. (1988)
Pilsbry, 1934

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Table 1 continued

Group/species First record Reference HB ST P/V

IP Galicia

Potamopyrgus antipodarum 1951 Boettger (1951) 1,5

(Gray, 1853)
1963 Lucas (1963) I U
Bulinus contortus 1845 Morelet (1845)
(Michaud, 1829)
1999 Rolán et al. (1999) U AQ
Pomacea canaliculata 2009 ** NR GEIB (2009) 1,2,3
(Lamarck, 1822)
Corbicula fluminea 1981 Mouthon (1981) 1,3,5
(Müller, 1774)
1989 Araujo et al. (1993) I FP
Dreissena polymorpha 1880 NR Servain (1880) 1,2
(Pallas, 1771)
Mytilopsis leucophaeta 1993 NR Garcı́a-Berthou et al. (2007) 1,2,5
(Conrad, 1831)
Xenostrobus securis 2005 2005 Garci et al. (2007) 5 I AO
(Lamarck, 1819)
Annelida
Branchiura sowerbyi 1970s NR Garcı́a-Berthou et al. (2007) 1,2,5
Beddard, 1892
Ficopomatus enigmaticus 1924 NR Garcı́a-Berthou et al. (2007) 5
(Fauvel, 1923)
Hirudo troctina (Johnson, 1893 1893 Blanchard (1893) 1,2,3 U U
1816)
Crustacea
Artemia franciscana 1980s NR Garcı́a-Berthou et al. (2007) 5
(Kellog, 1906)
Wlassicsia pannonica 1990s NR Garcı́a-Berthou et al. (2007) 4
Daday, 1904
Dolerocypris sinensis Sars, 1986 NR Garcı́a-Berthou et al. (2007) 3,4
1903
Ilyodromus viridulus 1996 NR Garcı́a-Berthou et al. (2007) 4
(Brady, 1886)
Cypris sp.(=Cypris 1986 NR Garcı́a-Berthou et al. (2007) 2,4
subglobosa Sowerby, 1840)
Isocypris beauchampi 1976 NR Garcı́a-Berthou et al. (2007) 2,4
(Paris, 1920)
Stenocypris major 1986 NR Garcı́a-Berthou et al. (2007) 3,4
(Baird, 1859)
Strandesia vavrai 1983 NR Garcı́a-Berthou et al. (2007) 4
(Müller, 1898)
Strandesia vinciguerrae 1986 NR Garcı́a-Berthou et al. (2007) 4
(Masi, 1905)
Tanycypris sp. 1988 NR Garcı́a-Berthou et al. (2007) 3,4
Acartia tonsa Dana, 1849 1990s NR Garcı́a-Berthou et al. (2007) 5
Synidotea laticauda 1996 NR Garcı́a-Berthou et al. (2007) 5
Benedict, 1897

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Table 1 continued

Group/species First record Reference HB ST P/V

IP Galicia

Cherax destructor Clark, 1983 NR Garcı́a-Berthou et al. (2007) 5

1936
Eriocheir sinensis Milne 1988 NR Garcı́a-Berthou et al. (2007) 5
Edwards, 1853
Pacifastacus leniusculus 1974 Carral et al. (1993) 1
(Dana, 1852)
2009 ** Xunta de Galicia (2009) U U
Palaemon macrodactylus 1999 NR Garcı́a-Berthou et al. (2007) 5
MJ Rathbun, 1902
Procambarus clarkii 1973 Garcı́a-Berthou et al. (2007) 1,2,4,5
(Girard, 1852)
1980s I IR
Rhithropanopeus harrisii 1990 NR Mariscal et al. (1991) 5
(Gould, 1841)
Insecta
Aedes albopictus (Skuse, 2004 NR Garcı́a-Berthou et al. (2007) 3
1895)
Stenopelmus rufinasus 2002 NR Fernández-Carrillo et al. (2005) 1,2,4,5
Gyllenhal, 1835
Trichocorixa verticalis 1997 NR Garcı́a-Berthou et al. (2007) 3,5
(Fieber, 1851)
Pisces
Abramis bjoerkna 1995 NR Elvira (1998) 1,2
(Linnaeus, 1758)
Abramis brama 2004 NR Garcı́a-Berthou et al. (2007) 2
(Linnaeus, 1758)
Acipenser baerii Brandt, 1995 NR Elvira and Almodóvar (1999)
1869
Alburnus alburnus 1992 NR Elvira (1995) 1,2
(Linnaeus, 1758)
Ameirus melas 1910 NR Elvira and Almodóvar (2001) 1,2
(Rafinesque, 1820)
Aphanius fasciatus 1997 NR Garcı́a-Berthou et al. (2007) 3
(Valenciennes, 1821)
Australoheros facetus 1943 NR Helling (1943) 1,2
(Jenyns, 1842)
Carassius auratus 17th 17th cent. Lobón-Cerviá et al. (1989) 1,2,3 N AT
(Linnaeus, 1758) cent.
Cobitis bilineata Canestrini, 2002 NR Garcı́a-Berthou et al. (2007) 1
1865
Cyprinus carpio Linnaeus, 17th Lozano-Rey (1935) 1,2
1758 cent.
18th cent. N AT
Esox lucius Linnaeus, 1758 1949 Gutiérrez-Calderón (1954) 1,2
2007** Press note (2008). C. H. Miño- U A
Sil

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Table 1 continued

Group/species First record Reference HB ST P/V

IP Galicia

Fundulus heteroclitus 1970 NR Garcı́a-Berthou et al. (2007) 4

(Linnaeus, 1766)
Gambusia holbrooki 1921 Nájera (1944) 1,2,3,4
(Girard, 1859)
[1940 N/I AT
Gobio lozanoi Doadrio & Authoct. 1998 Hervella and Caballero (1999) 1,2,3 N AB
Madeira, 2004
Hucho hucho (Linnaeus, 1968 NR Mondejar (1981) 1
1758)
Ictalurus punctatus 1995 NR Garcı́a-Berthou et al. (2007) 1,2,3
(Rafinesque, 1818)
Lepomis gibbosus 1910 Garcı́a-Berthou et al. (2007) 1,2
(Linnaeus, 1758)
2005 Comesaña and Ayres (2009) N AT
Misgurnus anguillicaudatus 2001 NR Franch et al. (2008) 3,4
(Cantor, 1842)
Micropterus salmoides 1952 Godinho and Ferreira (1994) 1,2
(Lacépède, 1802)
1982 Callejo et al. (1982) N A
Oncorhynchus mykiss 1898 Welcomme (1988) 1,2
(Walbaum, 1792)
1982 Callejo et al. (1982) NN AQ
Oncorhynchus kisutch 1983 NR Garcı́a-Berthou et al. (2007) 1
(Walbaum, 1792)
Perca fluviatilis Linnaeus, 1975 NR Garcı́a-Berthou et al. (2007) 1,2
1758
Phoxinus bigerri Kottelat, Authoct. Garcı́a-Berthou et al. (2007)
2007
1990s Hervella and Caballero (1999) N AB
Poecilia reticulata Peters, 2000 NR Garcı́a-Berthou et al. (2007) 1,4
1859
Pseudorasbora parva 1999 NR Caiola and de Sostoa (2002) 1
(Temminck & Schlegel
1846)
Rutilus rutilus (Linnaeus, 1910 NR Garcı́a-Berthou et al. (2007) 1,2
1758)
Salvelinus fontinalis 19th NR Lozano-Rey (1935) 1,2
(Mitchill, 1814) cent.
Sander lucioperca 1975 NR Garcı́a-Berthou et al. (2007) 1,2
(Linnaeus, 1758)
Scardinius 1910 NR Sostoa et al. (1990) 1,2
erythrophthalmus
(Linnaeus, 1758)
Silurus glanis Linnaeus, 1974 NR Garcı́a-Berthou et al. (2007) 1,2
1758
Tinca tinca Linnaeus, 1758 12th Garcı́a-Berthou et al. (2007) 2,3
cent.

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Biodivers Conserv (2010) 19:3471–3487 3483

Table 1 continued

Group/species First record Reference HB ST P/V

IP Galicia

\1991 Hervella and Caballero (1999) N A

Amphibia
Discoglossus pictus (Otth, 1900 NR Garcı́a-Berthou et al. (2007) 1
1837)
Reptilia
Trachemys scripta 1983 Tauxe et al. (1985) 1,2,3,4
(Schoepf, 1792)
1990s Galán (1999) I AT
Mauremys leprosa Authoct. 1,3
(Schweigger, 1812)
1990s Ayres and Garcı́a (2006) N AT
Mammalia
Neovison vison (Schreber, 1958 Bravo and Bueno (1992) 1,2,3,4
1777)
1960 Vidal-Figueroa and Delibes N/I FS
(1987)
Myocastor coypus (Molina, 1970 NR Saint-Girons (1973) 1
782)
Ondatra zibethicus 2004 NR Elosegui (2003) 1
(Linnaeus, 1766)
Azolla filiculoides Lam. 1920 Pereira et al. (2001) 1,2,3
1995 Romero-Buján et al. (2004) I AT
Eichhornia crassipes 1939 Moreira et al. (2005) 1,2,3
(Mart.)
2008** Press note (2009). N/I AT
C.H. Miño-Sil
Egeria densa Planch. 1995 Cirujano et al. (1995) 1,2,3
\2005 Pulgar and Izco (2005) I AT
Pistia stratiotes L. 1990 NR Domingues de Almeida 1,2,3
and Freitas (2006)
Elodea canadensis Michx. 1910 Domingues de Almeida 1,2,3
and Freitas (2006)
2006 Servia et al. (2006) N AT
Heteranthera limosa (SW.) 1995 NR Rodrı́guez et al. (1995) 3,4
Willd.
Heteranthera reniformis 1993 NR Zaragoza et al. (1993) 3,4
Ruiz & Pav.
Heteranthera rotundifolia 1996 NR Domingues de Almeida and 3,4
Griseb. Freitas (2006)
Myriophilllum aquaticum 1936 Domingues de Almeida and
(Velloso) Verdc. Freitas (2006)
1971 Laı́nz (1971) AT
Salvinia natans (L.) All. 1984 Bolós and Vigo (1984)
2005 Gómez Vigide et al. (2005) U

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3484 Biodivers Conserv (2010) 19:3471–3487

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