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Transcript
Reviewer #1:
Comments: The abstract is usually a short summary of the entire study. The present abstract is
only an introduction and leaves open the results and conclusions of the study. Of course there
are many results presented here, but some are more significant than others. The fact that Arion
belongs to the Arionidae family is not of crucial importance to be mentioned in an abstract.
Response: The abstract has been changed with taking into account comments from the
reviewers.
Comments: Drought resistance only for terrestrial species, not for Dreissena and other aquatic
species. Later in the paper is demonstrated that the species reproduced better with increased
humidity. It should be mentioned somewhere at the beginning in the introduction that for
decades the slug had been called Arion lusitanicus, and that many references recorded the
species under this name.
Response: This part has been supplemented with issues concerning about drought resistance.
It was added that A. vulgaris for many years had been called A. lusitanicus.
Comments: In the country list Spain is absent. I am not firm in the recent literature on this
species, but it should be mentioned in which year was the first reliably confirmed occurrence
of this species in Spain. France: the type locality of vulgaris is in France, so the date should be
1855, not 1955. A. vulgaris and A. rufus can be distinguished during the process of copulation.
This should be added. In the text it looks like a determination would only be possible by
anatomical dissection.
Response: All suggestions were taken into account and responses were added to the
manuscript. For example: “The origin distribution of this slug included Spain, Portugal and
Azores (Simroth 1891, Quick 1952, 1960; Regteren Altena 1971, Chevallier 1972). It looks like
A. vulgaris has expanded its range in many European countries over the last five decades. At
present, it occurs in France (1855), Great Britain (1952), Germany (1970), Slovenia (1970),
Italy (1971), Switzerland (1971), Austria (1972), Sweden (1975), Bulgaria (1983), Austria
(1984), Norway (1988), Belgium (1989), the Netherlands (1989), Finland (1990), Denmark
(1991), Poland (1993), Iceland (2003-2004), Greenland, Latvia, and Lithuania and has more
recently appeared on the Faroe Islands (Quick 1952, 1960, Ellis 1965, Schmid 1970, van
Regteren Altena 1971, Riedel & Wiktor 1974, Davies 1987, de Winter 1989, von Proschwitz
1992, 1994, von Proschwitz & Winge 1994, Wiktor 1996, Weidema 2006, Kozłowski 2007,
Slotsbo 2012)”, “When they co-occur in the same habitat, A. ater, A. rufus, and A. vulgaris can
only be identified by virtue of their genital morphology, process of copulation or by molecular
methods because the species look very similar (Quinteiro et al. 2005, Barr et al. 2009, Kałuski
et al. 2011, Slotsbo 2012). A. vulgaris has the following species-specific features: atrium small,
almost symmetrical, one-partite, bursa copulatrix oval; a fallopian tube with a short, thin
posterior end and a thick, rapidly expanding anterior end (Wiktor 2004). Copulation of A.
vulgaris may take from 4 to 5.5 hours (Kozłowski 2007), while in A. rufus this process takes 3
hours (Frömming 1954)”.
Comments: The sentence that A. vulgaris was first discovered by M-T in SW France should
have as reference MT 1855, not Anderson 2005. The type locality is uncertain, but likely, since
no types exist. Anderson 2005 could not determine a type locality without designating a type.
Who was Tylor? Van Regteren Altena 1955 misidentified a French species with A. lusitanicus.
As far as I know, A. vulgaris was not mentioned. Simply said, A. vulgaris was first discovered
by Van Regteren Altena in 1955, and in 1997 Castillejo discovered that a wrong name had been
used. The proposal to use A. vulgaris came in 2002 by Falkner et al. who had studied the French
literature. The statement that researchers used A. vulgaris once "again" is incorrect. A. vulgaris
1
had probably never been used before 2002/2005, because people actually did not know what it
was. Falkner et al. 2002 were the first to suggest using this name. The reference to lusitanicus
comes too late. This should be mentioned more at the beginning in the introduction.
Response: All comments were taken into account and manuscript was filled in with appropriate
information. The final version is: “A. vulgaris was first discovered in southwestern France in
1855 (Moquin-Tandon 1855). In 1907, Taylor claimed it was the same species as A. ater var.
rufa or A. subfuscus Draparnaud, 1805 (Taylor 1907). Then, in 1955, Regteren Altena said it
was actually Arion lusitanicus Mabille, 1868 (Castillejo unpublished). However, Castillejo
(1997) discovered that wrong name for this species has been used. A. lusitanicus, which was
described by Mabille in 1868, is common in central Portugal, and differs from the Spanish slugs
found throughout Europe in either external appearance or genital morphology. Molecular
analyses have confirmed that A. vulgaris and A. lusitanicus are actually two different species
(Quinteiro et al. 2005). These findings caused researchers to use the name A. vulgaris which
was proposed by Falkner et al. (2002). Consequently, in the older literature, when A. lusitanicus
is used to refer to an invasive species occurring in Europe, it is likely a synonym for A.
vulgaris”.
Comments: Which one is its native range? The statement needs a reference: In its invaded
range the species has since long invaded natural habitats, even in high altitudes in the Alps.
Response: It was added to the manuscript that “The origin distribution of this slug included
Spain, Portugal and Azores (Simroth 1891, Quick 1952, 1960; Regteren Altena 1971,
Chevallier 1972). It looks like A. vulgaris has expanded its range in many European countries
over the last five decades”. (line 55-58).
Comments: replace lusitanicus by vulgaris. Insert a return before "Another study", and give
the corresponding reference. It remains unclear what the times of activity should tell us. I would
expect data of other species to compare. Insert a return before "It is notable", because also here
we can compare between species.
Response: A. lusitanicus was replaced by A. vulgaris in manuscript. Comments have been
included in the final version of the text.
Comments: Arion fuscus: unclear which species is meant, the name seems to be controversial.
Maybe Arion subfuscus was meant. At the end of the paragraph it would be good to use the
terms robust and opportunistic to present the conclusions of that study.
Response: The paper Knop & Reusser 2012 was concerned about phenotypic plasticity of A.
lusitanicus compared with A. fuscus, not A. subfuscus. The conclusions were presented at the
end of this paragraph.
Comments: Ross et al. 2009: give the name of the species.
Response: Names of the species were added to the manuscript.
Comments: "presumed area of origin": which area was this exactly? How many samples had
been studied by Pfenninger et al. 2014? From which regions? How densely had the species been
sampled in France? What is the meaning of the 40 clades number? What is the meaning of the
average sequence divergence? How many species did they study? Add the country or region in
which the study was done. why is the story of expansion unusual? Do we actually know enough
about the story of expansion? It rather seems to me that we do not even know exactly the native
range from which the range expansion originated.
Response: This part has been more developed in manuscript.
2
Comments: It seems also that more finely tuned knowledge on this species is needed before
we can come to serious conclusions. This includes also conclusions on reasons and mechanisms
of range expansion. The current state of research is that A. vulgaris might possibly have had its
original range somewhere in Central Europe. Not exactly a very narrowly defined region, given
that the entire range of the species before the opening of the wall in 1990 had not been much
larger. Pfenninger et al. 2014 was only a beginning. It might be necessary to study 2 or 3 orders
of magnitude more samples in Europe than Pfenninger et al. 2014 did, and then see which
population groups seem to be related with which ones. In the same way as researchers found
out the original range of humans in some region in Africa.
Response: This part has been more developed in the manuscript.
Comments: Climate change is certainly a factor, but we also should keep in mind that slow
snails and slugs may still invade central and northern European regions as a result of an reinvasion after the ice age. The average size of species in Sweden is still smaller than in S
Europe, because minute species are faster in natural range expansions. By logical consequence
one must assume that there are still places free in Sweden to provide habitats for larger
terrestrial mollusc species. This effect should be ruled out in the analysis of invasive species.
Are there species among those which reach central and northern Europe with artificial help that
reach the area just faster that they would have reached the same region somewhere in the future
by natural means? And from which regions could slow species invade colder regions in Europe,
if not from warmer regions? There are no cold regions which host so many species that they
can provide many invaders. Also statistically this should be evaluated. The warm regions
naturally host more species, so potentially they can provide many more species to invade other
regions, colder regions than colder regions could provide to invade warmer regions.
Response: This topic will be more developed in our future studies connected with the project
which we are realizing. These papers will be connected with climate changes and its impact on
biodiversity and spread of invasive species.
3
Reviewer #2:
Comments: Generally, the manuscript looks like it has been done quite carelessly. Even the
structure of sections is a bit puzzling. The only main sections (according to the journal style)
are Abstract, Introduction and References. Between the latter two are additional, small
headlines (one of them fat): Traits enhancing competitivity, Greater phenotypic plasticity,
Enemy release hypothesis, Rapid response to natural selection, The case of Arion vulgaris,
Climate change, Conclusions and future directions. So these are the main chapters dealing with
questions for the slug’s success - there should be a headline for this. Considering the task of the
manuscript (as I understand it), the manuscript consists of two parts (not clearly separated). The
first section (headed “Introduction”) provides the bulk of the general review about biology, pest
status and spread. The rest deals mainly, but not exclusively, with potential reasons for the great
success of the species.
Response: The manuscript was reorganized and divided into several sections, such as: Abstract,
Introduction, Naming and distribution, Differentiation of species, Gastropod pests control
method, Biology of A. vulgaris, Potential reasons for the great success of Arion vulgaris, Traits
enhancing competitivity, Phenotypic plasticity, Enemy release hypothesis, Rapid response to
natural selection, Climate change, The case of Arion vulgaris, Conclusions and future
directions, Acknowledgements, References. The manuscript has been more developed.
Comments: As an introduction (in case I misunderstood and the authors do not aim for a
general review), this first part is far too long. As a review it is quite poor. The references look
like an arbitrary choice of articles, and the authors have rarely put effort into tracing back
original sources. In some cases I doubt the authors have really read the references thoroughly.
Also problematic are statements presented as facts that are based on online websites that
themselves do not provide proper references. Such websites are convenient sources, but this
does not fulfill the expectations put into a review and merely adds to the pile of unproven
statesments which, by pure repetition, become accepted as facts. A few examples (this is not
comprehensive - there are more such cases): 1. page 3, second paragraph: “It has also newly
arrived in Romania and Slovenia (Laznik et al. 2010)”. The cited article is about nematode
infection; A. vulgaris specimens (lots) used for the laboratory tests were collected at two areas
in Slovenia. No mention about its first discovery or spread in Slovenia, zero about Romania. In
fact, A. vulgaris had already been found in Slovenia in 1970 (see Wiktor 1996 or a review about
the slugs of Slovenia by Vaupotič & Velkovrh 2002). There is a reference available about
Romania (Papureanu et al. 2014 in Malacologica Bohemoslovaca 13: 6-11).
Response: This part has been more developed. Introduction was divided into smaller parts such
as Introduction, Naming and distribution, Differentiation of species, Gastropod pests control
method. The final version of this part is as follows: “The native range of A. vulgaris is not
known for certain. For many years, it was believed that the slug came from southwestern Europe
(Roth et al. 2012) and began its spread over 40 years ago on the Iberian Peninsula. It looks like
A. vulgaris has expanded its range in many European countries over the last five decades. At
present, it occurs in Great Britain (1952), France (1955), Germany (1970), Italy (1971),
Switzerland (1971), Austria (1972), Sweden (1975), Bulgaria (1983), Austria (1984), Norway
(1988), Belgium (1989), the Netherlands (1989), Finland (1990), Denmark (1991), Poland
(1993), Iceland (2003-2004), Greenland, Latvia, and Lithuania and has more recently appeared
on the Faroe Islands (Weidema 2006, Kozłowski 2007, Slotsbo 2012). It has also newly arrived
in Romania and Slovenia (Laznik et al. 2010). A. vulgaris has been present in USA since 1998
(Rabitsch 2006, Barr et al. 2009). However, some recent findings contradict the idea of an
Iberian origin. Pfenninger et al. (2014) suggested that it is highly probable that the species is
actually native to central Europe: phylogeographic analyses using mitochondrial (COI) and
nuclear (ZF) markers combined with species distribution modelling found that no haplotypes
4
from Spain or western France co-occurred with haplotypes from central Europe. However, this
discovery aside, A. vulgaris clearly exhibits invasive features in the countries where it has most
recently appeared”.
Comments: “A. vulgaris has been present in USA since 1998 (Rabitsch 2006, Barr et al.
2009).” In fact, the article by Barr et al. states that “the A. ater and A. vulgaris morphologies
not yet detected in USA”. The reference by Rabitsch 2006 is the DAISIE Factsheet about A.
lusitanicus and does itself not provide a reference for this. In fact, it is based on an unconfirmed
online record and should be omitted.
Response: This part has been corrected and changed. The reference Rabitsch 2006 was omitted.
Sentence “A. vulgaris has been present in USA since 1998 (Rabitsch 2006, Barr et al. 2009)”
was deleted.
Comments: “Castillejo (1997) discovered that A. lusitanicus ... is endemic to Serra Arrábida,
Portugal, ...” In fact, the cited reference does not mention endemism, but reports that this species
is common in central Portugal, and that other records from the Iberian Peninsula require
reinvestigation.
Response: This part has been changed to “However, Castillejo (1997) discovered that A.
lusitanicus, which was described by Mabille in 1868, is very common species in Portugal, and
differs from the Spanish slugs found throughout Europe in either external appearance or genital
morphology”.
Comments: The species-specific features provided for the genital anatomy imply that the
authors are not really familar with the morphology. The “a small atrium; a onepiece, almost
symmetrical, oval spermatheca” is an unfortunate (google?) translation of a reference in Polish
(Wiktor 2004, which itself is not the best choice). The correct translation is: “Atrium small (!),
almost symmetrical, onepartite. Bursa copulatrix oval.” So, the atrium is symmetrical and onepartite, not the bursa. Lots of Arion species have an oval bursa copulatrix, and they are all onepartite and more or less symmetrical. In fact, the main diagnostic characters are the small atrium
in combination with the big distal free-oviduct and the shape of the ligula.
Response: This part has been changed to “Monitoring the spread of A. vulgaris is challenging
because the species is hard to distinguish from the other closely related, large arionids (A. ater
L. 1758; A. rufus L. 1758; A. vulgaris; A. magnus Torres Minguez 1923; A. lusitanicus Mabille
1868; A. flagellus Collinge 1893) that occur in Europe. When they co-occur in the same habitat,
A. ater, A. rufus, and A. vulgaris can only be identified by virtue of their genital morphology,
process of copulation or by molecular methods because the species look very similar (Quinteiro
et al. 2005, Barr et al. 2009, Kałuski et al. 2011, Slotsbo 2012). A. vulgaris has the following
species-specific features: atrium small, almost symmetrical, one-partite, bursa copulatrix oval;
a fallopian tube with a short, thin posterior end and a thick, rapidly expanding anterior end
(Wiktor 2004). Copulation of A. vulgaris may take from 4 to 5.5 hours (Kozłowski 2007), while
in A. rufus this process takes 3 hours (Frömming 1954). To complicate matters further, A. ater
and A. rufus can hybridise, and there is some evidence that they hybridise with A. vulgaris as
well (Evans 1986, Roth et al. 2012, Dreijers et al. 2013)”.
Comments: “Spanish slugs can excrete ingested toxins in their muscus, which might serve as
a defence mechanism (Aguiar & Wink 2005).” I can well imagine that the species is able to do
this, but the reference is inappropriate: The authors of the cited reference injected the toxin
dorsally through the skin, and the main bulk of excretion was within 10 minutes after injection.
So, this does not prove that the slugs can excrete in their body mucus toxins taken up with food.
5
Such incorrect citations add to the confusion and pool of unproven statements, and repetitions
of them tend to make them appear true. This is not what I would expect from a review.
Response: This part has been deleted from the manuscript.
Comments: On the second part (reasons for the success). While this is an interesting question,
the manuscript does not exploit this topic exhaustively. Neither do I find the different options
(hypotheses) explored sufficiently.
Response: This part of the manuscript has been more developed.
Comments: “It is thought that A. vulgaris outcompetes the autochthonous A. ater and A. rufus.”
There are quite a number of references available, but only one is cited.
Response: Another references were added such as von Proschwitz 1997, Kappes & Kobialka
2009, Roth et al. 2012, Dreijers et al. 2013, Hatteland et al. 2013.
Comments: Much of the second paragraph (“Another study found ... similar to what is seen in
A. ater (Grimm & Paill 2001).”) seems irrelevant. It deals with activity patterns of A. vulgaris
without any obvious relation to its competitiveness or differentiation from other species. This
would seem better placed in the general review part which deals already with the influence of
photoperiod.
Response: This part was deleted and transferred to another paragraph.
Comments: Greater phenotypic plasticity. The only relevant reference seems to be the study
by Knop & Reusser (2012): a comparison of A. vulgaris with A. fuscus. However, I would have
wished a somewhat careful interpretation and the comment that these two species are rather
different anyway. Why not a comparison with the closer relatives A. ater or A. rufus?
Response: In the paper Knop & Reusser (2012) authors focused on phenotypic plasticity of
invasive slug, A. vulgaris, compared with results obtained for native slug, A. fuscus. The main
aim of this study was verification which scenario is suitable for A. vulgaris in the context of
how invader may benefit from phenotypic plasticity. We claimed that these species are different
in terms of their status (invasive vs. native).
Comments: Observations showing that juveniles start growing faster at higher temperature are
not very surprising for a mollusc. At least, I am missing an explanation why this should reflect
a “greater phenotypic plasticity” -- greater than what? Again, this seems irrelevant here and fit
better in the general review.
Response: Some parts has been deleted, also word greater.
Comments: Enemy release hypothesis. There is no clear evidence mentioned that A. lusitanicus
might have less enemies than related native species or than in its native area. The fact that
intensely farmed areas may have less predators would hold for native as well as invaded areas
and also for native species.
Response: This part has been more developed in the manuscript. Final version of this paragraph
is as follows: “The enemy release hypothesis (ERH) states that the lack of natural enemies in
an invader’s introduced range influences its abundance or impact (e.g., estimated using
individual size, population abundance, or propensity to displace native species) (Lee 2002,
Torchin et al. 2003, Colautti et al. 2004). When Ross et al. (2009) compared the occurrence of
parasites in a slug species in its native range in the UK and its introduced range in the USA,
they found support for the ERH (Ross et al. 2009). However, the situation may be a bit more
complex. Colautti et al. (2004) suggested that there are strong, enemy-specific effects on host
survival and that hosts have developed tailored defences. It is the release from the specific
6
enemy that causes direct changes to survivorship, fecundity, biomass, or demographic variables
that matters. Indeed, it is of little consequence if a species escapes an enemy or enemies against
which it is well defended (Colautti et al. 2004). In a study of carabid beetle predation on A.
vulgaris in Norway, Hatteland (2010) found that the slug still faces natural enemies even in its
introduced range. It was observed that Pterostichus niger, P. melanarius, Carabus nemoralis,
C. violaceus and the staphylinid beetle Staphylinus erythropterus to prey on eggs and newly
hatched slugs of A. vulgaris (Pail 2000, Paill et al. 2002, Hatteland et al. 2010, Hatteland et al.
2011). Pterostichus species are more restricted in the size of prey taken compared to larger and
more specialised predators, such as Carabus spp. (Hatteland et al. 2010). C. nemoralis was
shown to be a potentially important predator of the alien A. vulgaris in spring and may
contribute to conservation biological control (Hatteland et al. 2011). This species killed and
consumed juvenile slugs of up to 1 gram (Hatteland 2010). However, in intensively farmed
areas, predator populations may be reduced in size, leading to a greater abundance of A.
vulgaris. Temperature can also affect rates of predation on slugs. The activity threshold of C.
nemoralis has been found to be 4°C, and activity is greater when temperatures rise in spring,
while activity is not correlated with temperature later in the season (Hatteland et al. 2011).
Comments: The only enemies listed are from a single reference (Hatteland, 2010). But there
are certainly more references available.
Response: The other publications have been added to this part and all paragraph has been
reorganised and more developed. The final version is: “The enemy release hypothesis (ERH)
states that the lack of natural enemies in an invader’s introduced range influences its abundance
or impact (e.g., estimated using individual size, population abundance, or propensity to displace
native species) (Lee 2002, Torchin et al. 2003, Colautti et al. 2004). When Ross et al. (2009)
compared the occurrence of parasites in a slug species in its native range in the UK and its
introduced range in the USA, they found support for the ERH (ROSS et al. 2009). However, the
situation may be a bit more complex. Colautti et al. (2004) suggested that there are strong,
enemy-specific effects on host survival and that hosts have developed tailored defences. It is
the release from the specific enemy that causes direct changes to survivorship, fecundity,
biomass, or demographic variables that matters. Indeed, it is of little consequence if a species
escapes an enemy or enemies against which it is well defended (Colautti et al. 2004). In a study
of carabid beetle predation on A. vulgaris in Norway, Hatteland (2010) found that the slug still
faces natural enemies even in its introduced range. It was observed that Pterostichus niger, P.
melanarius, Carabus nemoralis, C. violaceus and the staphylinid beetle Staphylinus
erythropterus to prey on eggs and newly hatched slugs of A. vulgaris (Pail 2000, Paill et al.
2002, Hatteland et al. 2010, Hatteland et al. 2011). Pterostichus species are more restricted in
the size of prey taken compared to larger and more specialised predators, such as Carabus spp.
(Hatteland et al. 2010). C. nemoralis was shown to be a potentially important predator of the
alien A. vulgaris in spring and may contribute to conservation biological control (Hatteland et
al. 2011). This species killed and consumed juvenile slugs of up to 1 gram (Hatteland 2010).
However, in intensively farmed areas, predator populations may be reduced in size, leading to
a greater abundance of A. vulgaris. Temperature can also affect rates of predation on slugs. The
activity threshold of C. nemoralis has been found to be 4°C, and activity is greater when
temperatures rise in spring, while activity is not correlated with temperature later in the season
(Hatteland et al. 2011)”.
Comments: Rapid response to natural selection. Not a single piece of evidence or reference for
A. vulgaris is provided.
Response: This part has been developed and two paragraphs has been added: “Stamps (2007)
argued that selection for high individual growth rates would increase mean levels of risk-taking
7
behavior across populations. Similarly, a high population growth rate may further encourage
dispersal. The high life-time productivity of A. vulgaris with over 400 eggs laid by a single
individual may be a strategy to compensate for a higher mortality and at the same time allow
rapid population growth in newly colonized locations (KAPPES et al. 2012). It was also
confirmed that A. vulgaris is less sensitive to otherwise aversive stimuli, and thus more likely
to utilize novel environments and otherwise unusual dispersal routes (Slotsbo et al. 2011,
Kappes et al. 2012). It is expected that species, in which selection is against individuals taking
dispersal-related risks under undisturbed conditions, may acquire (or loose) traits and become
successful invaders under changing environmental conditions. A. vulgaris it is suitable
organisms for testing this hypothesis and dispersal behavioral syndromes, which, according to
Cote et al. (2010), include traits such as locomotor and feeding activity, boldness, exploration,
sociability and aggressiveness (Kappes et al. 2012)”.
Comments: The case of Arion vulgaris. Here (or in the conclusions), one would expect a kind
of summary of which data about the slug’s biology are relevant for the different hypotheses,
but I cannot find this here or anywhere else. Instead, the authors mainly and extensively refer
to a recent publication by Pfenninger et al. (2014) claiming to have “found that” A. vulgaris is
native to central Europe. Considering the narrow sampling of the cited study, I would have
wished an at least slightly careful discussion. And: if the authors are so much in favour of the
study, shouldn’t they be particularly critical with statements that the species has been unusual
in its rapid spread? The vast majority of previous studies dealing with the spread and success
of A. vulgaris (and cited in this manuscript) are based on the assumption that the origin of A.
vulgaris is somewhere in SW Europe.
Response: This part of the manuscript has been more developed and corrected.
Comments: “It has also been suggested that hybridisation has resulted in A. rufus disappearing
in areas invaded by A. vulgaris (Dreijers 2013; Allgaier 2015).” In fact, Allgaier is very
sceptical about this, but this is not mentioned in the manuscript.
Response: It was added to the manuscript that “However, study conducted by Allgaier (2015)
did not confirmed hybridisation between A. lusitanicus and A. rufus in the field. Different
habitats and overlapping of these two species may contribute to this (Allgaier 2015).
Comments: Climate change. It seems reasonable to assume that A. vulgaris may profit from
global warming, but beside a single reference about observations in SE Poland, no further
relevant data on A. vulgaris are presented. This seems a bit weak, the more as this weakness is
not discussed.
Response: This chapter has been more developed and it was added to this part that “In Norway,
using geoclimatic parameters such as average monthly temperature and precipitation, described
a potential future distribution of A. vulgaris in this country. The conducted study revealed that
large parts of coastal and lowland Norway will be potentially suitable areas for this pest. The
results showed that A. vulgaris is clearly favoured by the mild and wet Atlantic climate of the
western coast. The damage reported in gardens and horticulture (e.g. strawberries) related to
this pest has been especially pronounced in coastal areas. It can be expected that habitat
structure and its resulting micro-climate may be more important for the establishment of A.
vulgaris populations. Moreover, regardless of regional differences, precipitation throughout the
whole country is not a limiting factor for A. vulgaris (Hatteland et al. 2013)”.
Comments: Conclusions and future directions. I could not spot any conclusions except the need
for more studies.
Response: As it was presented above, all manuscript has been more developed and corrected.
8