Common juniper, an overlooked conifer with high invasion potential in protected areas of patagonia

ABSTRACT The benefits of early detection of biological invasions are widely recognized, especially for protected areas (PAs). However, research on incipient invasive plant species is scarce

compared to species with a recognized history of invasion. Here, we characterized the invasion status of the non-native conifer _Juniperus communis_ in PAs and interface areas of Andean

Patagonia, Argentina. We mapped its distribution and described both the invasion and the environments this species inhabits through field studies, a literature review, and a citizen science

initiative. We also modeled the species’ potential distribution by comparing the climatic characteristics of its native range with those of the introduced ranges studied. The results show

that _J. communis_ is now widely distributed in the region, occurring naturally in diverse habitats, and frequently within and close to PAs. This species can be considered an incipient

invader with a high potential for expansion in its regional distribution range, largely due to its high reproductive potential and the high habitat suitability of this environment. Early

detection of a plant invasion affords a valuable opportunity to inform citizens of the potential risks to high conservation value ecosystems before the invader is perceived as a natural

component of the landscape. SIMILAR CONTENT BEING VIEWED BY OTHERS EARLY WARNING OF TWO EMERGING PLANT INVADERS IN EUROPE Article Open access 05 April 2025 INVASION DYNAMICS OF THE EUROPEAN

BUMBLEBEE _BOMBUS TERRESTRIS_ IN THE SOUTHERN PART OF SOUTH AMERICA Article Open access 27 July 2021 INVASIVE ALIEN SPECIES OF POLICY CONCERNS SHOW WIDESPREAD PATTERNS OF INVASION AND

POTENTIAL PRESSURE ACROSS EUROPEAN ECOSYSTEMS Article Open access 19 May 2023 INTRODUCTION Protected areas (PAs) worldwide are recognized as a key component of the broad response to the

environmental degradation caused by global change, mainly because of their crucial role in conserving biodiversity1,2. Paradoxically, these areas are suffering increased degradation due to

processes related to global change, biological invasions being one of the most important drivers associated with this phenomenon3,4,5. In particular, the biodiversity and integrity of

several PAs around the world are being jeopardized by the invasion of introduced plants1, a process fostered especially by human activity6,7. Since PAs are not entirely excluded from the

major threats to biodiversity, the unique biological reservoirs contained in these areas are being increasingly compromised. Most PAs are interspersed with or adjacent to a mosaic of

landscapes altered by human influence3,8. The spatial configuration of these landscapes can facilitate a network of potential pathways for introduced species9. Indeed, the abundance and

composition of non-native plant species in PAs are strongly influenced by their surroundings, mainly due to the rapid colonization of these species from belt zones10. Non-native plant

species are undesirable in PAs, and those which are invasive are considered a priority for research and management10,11,12. In this regard there is abundant research on plant species with a

recognized history of invasion and conspicuous impacts on natural areas (e.g. ref.13,14; however, research on incipient invasive species is relatively scarce (e.g. ref.15 and references

therein). This is despite the ecological knowledge that incipient plant invaders may respond to efficient management strategies before they advance in the invasion process16 and have a

significant ecological impact, at which point their eradication becomes unlikely17. Although the benefits of early detection of incipient invasion in natural habitats are well recognized, so

are the difficulties associated with it18,19,20. Detection of early invasion foci is usually fortuitous16, and citizen collaboration is important in increasing the probability of

registering these situations. Public engagement is being enhanced by collaborative projects, led by professional scientists, that seek to compile information on potentially invasive

species20,21. In particular, citizen science has emerged as a powerful tool for detecting and mapping the distribution of recent invasive species and obtaining diverse bio-ecological

information on them20,22,23. This knowledge can provide insights into the invasion stage, the mechanisms behind the invasion, and the invader’s potential ecological impact, which can be

context-dependent24,25. Climate is recognized as the single most important factor determining the distribution of plant species at a large scale26,27. Thus, a frequently used approach to

predict where a species might invade is analysis of the climatic similarity between its native range and areas outside it28,29, even for plant species with no invasive history30. This

approach has been used for invasion risk assessment of non-native conifer species in areas of their introduced ranges throughout the Southern Hemisphere28,30, where they pose a significant

threat to the diversity and functioning of native ecosystems31 and even PAs32,33. In particular, climate matching can be a valuable tool for estimating suitable areas for potentially

damaging non-native conifers with incipient invasion. By cross-referencing information, it is possible to prioritize the search for and control of new invasion foci in, for example, PAs with

high invasion risk. The PAs of Andean Patagonia are no exception in terms of their high vulnerability to an increasing number of introduced plant species34,35. In this region, increasing

anthropogenic pressure on PAs acts as a catalyst for new invasions of introduced plant species whose invasive status, ecology, impact, and distribution are mostly unknown. This is

exemplified by the conifer _Juniperus communis_ L. (native to temperate regions of the boreal hemisphere), which has been identified as a potential high-risk invader of climatically suitable

areas in Africa30 and Oceania28. At the southernmost tip of South America, Argentina, _J. communis_ has recently been officially cataloged as an invasive species (Ministerio de Ambiente y

Desarrollo Sostenible 2021). Despite this, no studies have addressed its invasion status, distribution, or potential expansion range, especially in the areas of the country where it may

represent a risk to biodiversity, such as the PAs. This species already has three validated records in PAs of Andean Patagonia, according to the Biodiversity Information System which

provides biological information on the species, and PAs of Argentina (www.sib.gob.ar). However, _J. communis_ can be frequently seen in PAs of Andean Patagonia, which suggests that it is

under-recorded, probably because of its incipient invasion (i.e. the earliest stage of the invasion process). This assumption of an incipient invasion is supported by the lack of _J.

communis_ registers in key reference literature describing the regional flora36, including literature focusing on introduced plant species in the main PAs of the region35,37. In addition,

this species is increasingly valued as the raw material for producing gin, an alcoholic beverage that is booming internationally. This encourages its cultivation in the area, which can

increase the source of propagules for invasions in nearby PAs. It can also be seen in gardens; however, its incidence as an ornamental plant, and therefore the importance of this type of use

as a source of propagules, is as yet unknown. Here, we characterized the invasion status of the non-native conifer _J. communis_ in PAs and interface areas of Andean Patagonia, Argentina,

by mapping its distribution and describing both the invasion and the environments this species inhabits. We registered the type of invaded habitats, species abundance, its spatial

configuration pattern, the accompanying woody species, the species’ reproductive potential (i.e. presence of reproductive plants and seedlings), its importance as an ornamental plant, and

its occurrence in PAs and associated areas. We also modeled the potential distribution of the species by comparing the climatic conditions in its introduced range in Patagonia with those of

its native distribution range. We used different methodological approaches to acquire data on the species in the region: a literature search, field sampling, and citizen collaboration. To

our knowledge, this is the first work to provide information on _J. communis_ as an invader of a South American country. We address key descriptive aspects of the current _J. communis_

invasion that provide clues to the ecological mechanisms involved in its spread. Knowledge of the potential distribution of _J. communis_ could be useful in determining the invasion risk the

species presents for high conservation value ecosystems of Patagonia. RESULTS _JUNIPERUS COMMUNIS _IN ANDEAN PATAGONIA We compiled 174 occurrences of _J. communis_ in the region (58.6% from

field sampling, 33.9% from citizen contributions, and 7.4% from the literature review); > 90% of these records were from PAs (Fig. 1). We detected the presence of _J. communis_ within

eight PAs and close to another seven in the region (Table 1). Almost 100% of occurrences (sampled or reported) were associated with disturbed environments, mostly represented by roadsides

(gravel or paved) and trails (Fig. 2). Field sampling data indicated that _J. communis_ was found most frequently in forests (48%), followed by shrublands (26%), with the lowest

representation in steppe environments (5%; Fig. 2). We found an equal occurrence of the species in natural and urban habitats, with only a small percentage of ornamental use (Figs. 2, 3).

Regarding the spatial distribution pattern, _J. communis_ was frequently found as isolated individuals (62%), followed by thickets (21%) and, to a lesser extent, both patterns in the same

site (5%; Figs. 2, 3). The species was found mostly at low abundance (2–10 individuals in 45% of the occurrences), followed by a single individual (24%), medium density (11–100 individuals

in 16% of the occurrences), and high density (> 100 individuals in 11% of the occurrences; Figs. 2, 3). Fruited individuals and seedlings were observed in ca. 70%, and 50% of the

registers, respectively (Fig. 3), which could represent an underestimation of seedling presence since the understory of some sites was difficult to explore due to dense vegetation. In

addition, we registered 24 main woody species accompanying _J. communis_, half of which were native (Fig. 4). The most frequently found native species were _Austrocedrus chilensis_

(Cupressaceae), _Maytenus boaria_ (Celastraceae), _Nothofagus dombeyi_ (Nothofagaceae), and _Lomatia hirsuta_ (Proteaceae). Among the non-native species the most frequently found were _Pinus

contorta_ (Pinaceae), _Rosa rubiginosa_ (Rosaceae), and _Cytisus scoparius_ (Fabaceae) (Figs. 1, 4). The literature review afforded 18 records of _J. communis_ cited as a naturally

established species in the Andean Patagonian region (Neuquén, Río Negro, and Chubut provinces), with the oldest record dating back to 2002 (Table 2). Most of the records corresponded to PAs

(82%), including four national parks. In most of the studies the inclusion of _J. communis_ was not intentional, rather it appeared when describing vegetation, or when listing introduced

species (Table 2). Only three studies considered the species as the focus of their research, although none of these recognized it as invasive (Table 2). POTENTIAL DISTRIBUTION AND

BIOCLIMATIC MATCHING The Andean Patagonian region showed a highly climatically suitable land area for _J. communis_ occurrence (Figs. 1, 5), with the area of the best fitting model covering

all major PAs in the region (Fig. 1). The area of greatest suitability occupies the region near the Andes, from central to southern Argentina, becoming longitudinally wider towards the north

of Andean Patagonia and extending eastward into the southern part of Río Negro and northern Chubut. The three environmental variables of major importance to the _J. communis_ distribution

model were the mean temperature of the warmest and coldest quarters and the precipitation of the coldest quarter (Table S1). The functional relationship between the four continuous predictor

variables studied and the predicted habitat suitability (Fig. S1), shows that the highest values for probability of presence are given for a mean temperature of the warmest quarter between

10 and 20 °C, a mean temperature of the coldest quarter between − 20 and 10 °C and precipitation of the coldest quarter superior to 200 mm (Fig. S1). The bioclimatic variables analyzed for

the PAs of Andean Patagonia differed from those of the native range of _J. communis_: PAs in Andean Patagonia showed higher mean temperatures and precipitation in the coldest quarter and

lower mean temperature and precipitation in the warmest quarter than the species’ native range (Fig. 5). These results were supported by the jackknife test (Fig. S2), which also showed that

the mean temperature of the coldest quarter had the most useful information when considered alone (highest gain in isolation), and information that was not present in the other variables

(highest gain decrease when omitted). DISCUSSION _Juniperus communis_ has been in the Andean Patagonian region for at least 90 years, and the results of our work show that the time lag

between its introduction and its invasion is coming to an end. Records of an intentional introduction of this species in northwestern Patagonia date back to the 1930s, when it was to be

cultivated for ornamental purposes55. Seventy years later the species was registered in the same location, but occurring naturally56. Our results indicate that _J. communis_ has achieved a

wide distribution in Andean Patagonia, occurring naturally in diverse habitats, with numerous occurrences inside and close to PAs. The information we gather in this study allows us to

characterize _J. communis_ as an incipient invader with high potential for expansion of its regional distribution range. The likelihood of this spread can be largely determined by the high

reproductive potential of the species and high habitat suitability of the invaded region. The wide distribution of _J. communis_ in the region can be partially explained by seed dispersal,

which probably occurs via endozoochory by common species of the regional fauna. The main dispersers of _J. communis_ in its native range are birds of the genus _Turdus_, which is also

represented in Andean Patagonia. In other regions, for example, in England, _T. viscivorus_, _T. merula_, and _T. philomelos_ have been identified as the main dispersers of _J. communis_

seeds57, while in mountainous regions of the European Mediterranean the seeds of this species are dispersed almost exclusively by _T. torquatus_ and _T. viscivorus_58,59. In Andean

Patagonia, the genus _Turdus_ is mainly represented by _T. falcklandii_ (www.sib.gob.ar)60, which consumes fleshy fruits of common shrub species61,62, including those of _J. communis_

(Lambertucci S., pers. comm.). Additionally, there are records of apparently viable seeds of this species in the feces of hares63 and red deer (Relva A., pers. comm.). It is interesting to

note that _T. falcklandii_ is the most important frugivore present during winter61. Unlike most functionally equivalent native woody species, during winter _J. communis_ bears fruits, which

may represent a reproductive advantage for the invader64. The preliminary evidence described for this incipient invader highlights the importance of studying aspects of its reproductive

ecology (e.g. phenology) that may provide clues to mechanisms that facilitate its spread and its potential impact on the recipient communities. Although dispersal is an important factor in

determining plant species’ spread, climatic conditions are decisive in determining establishment success. The results showed high habitat suitability for the species based on climate

variables, which was evidenced in the high proportion of the sampled sites with seedlings and fruit-bearing individuals; that is, plant stages indicative of population growth. This contrasts

with what is currently happening to _J. communis_ in different areas of its native range, where the number of populations and their size have decreased drastically, mainly due to a lack of

natural recruitment65,66. This is mainly associated with a low percentage of viable seeds67, which could be due in part to climate effects. For example, Garcia et al.65 demonstrated that

rainy spring periods (short but heavy storms) in open shrublands of the Mediterranean mountains negatively affected the viability of _J. communis_ seeds by impeding pollen dispersal. This

effect is probably not as pronounced in the areas of incipient invasion in Patagonia, where rainfall is scarce during the period of pollen dispersal (i.e. the warmest quarter of the year)

and is even lower than that registered in the species’ native range (Fig. 5). Furthermore, in Patagonia, the scarcity of rainfall during the warmest period of the year is expected to be

accentuated by climate change in the coming years68,69. The disparities in climatic variables between native and introduced ranges could indicate adaptation70 or phenotypic plasticity71.

This highlights the importance of closely monitoring species like _J. communis_, to evaluate a potential climatic niche shift72,73, and to reassess these invasions in prospective climate

change scenarios. Many other effects may be involved in _J. communis_ pollination failure in particular57,74, and in its population decline in general in areas of its native range57,67.

Concerning the latter, the incidence of a recently invasive pathogen in Europe, the oomycete _Phytophthora austrocedri_, is causing widespread mortality in _J. communis_ native

populations75. This pathogen is already present in Andean Patagonia76,77, so it would be interesting to evaluate its incidence in the introduced populations of the woody invader, as well as

the incidence of other factors that negatively affect the species in its native range but seem not to hinder its expansion in Patagonia. Belt zones control the number of invaders in PAs,

determining the entry and spread of these species into the natural vegetation matrix7,9. _Juniperus communis_ was associated with roads and walking trails that were in close contact with

natural vegetation, which was evidenced by the high proportion of native woody species that accompanied it_._ Among the accompanying species were trees characteristic of Andean-Patagonian

forests, such as _Austrocedrus chilensis_ and _Nothofagus dombeyi_. In turn, the high frequency of these tree species reflects the prevalence of _J. communis_ in forest ecosystems. Among the

most frequently cited non-native species there were long-time invaders, highly adapted to human-modified environments, such as _Pinus contorta_ and _Rosa rubiginosa_ (Fig. 4;

www.sib.gob.ar). Considering that forest habitats are suitable for _J. communis_ invasion, and that disturbed areas represent expansion opportunities for this species, the increased

degradation caused by new trails (that deviate from those officially delimited) produced by domestic animals and visitors to PAs78 is of great concern. On the other hand, although medium to

high abundance invasion was observed in ca. one-fourth of the sampled sites, it was common to find individuals in small groups or alone (i.e. isolated from other conspecifics but not from

other woody species). While individuals established far from parent plants may indicate an increase in the spatial occupancy of the species, it also reveals that conditions for its control

in areas of conservation concern may be favorable (i.e. small population size15,79). As pointed out for other conifer invasive species80, the relatively low growth rate of woody plants

affords a time window during which on-the-ground action can be taken before the incipient invasion takes hold – even a single plant can constitute a significant propagule source to the

surroundings15. Unlike other woody plants that became invaders of natural environments associated with urban areas81,82, _J. communis_ was infrequently found as an ornamental or a living

fence plant. Therefore, the current use of this species does not represent a major threat in terms of invasion spread. However, attention should be paid to other human-induced propagule

sources. In light of the increasing valuation of this species for gin production, it would be interesting to investigate the importance of emerging cropping areas as a source of propagules

that could spread to natural areas, as well as the generation of protocols to minimize its potential dispersal and consequent invasion risk. On the other hand, while fruit harvesting by

local people from natural populations can reduce propagule pressure, it also favors positive public perception of the species as being of value as an economic resource83. Thus, the control

of incipient invaders could be a particularly difficult challenge in areas associated with PAs, due to the cultural importance and economic value certain invasive species can represent for

residents and visitors84,85. CONCLUSIONS We present here the first documentation of the distribution and descriptive characteristics of an incipient invasion of _J. communis_ in PAs of

Andean Patagonia, Argentina. Although the results indicate that the species has high spreading potential, they also show that this is an opportune moment for its control in areas that merit

conservation. Since belt areas are important in mediating introduced plant biodiversity in PAs, raising citizen awareness of environmental issues such as plant invasion is crucial. Citizen

science is a powerful tool when used as a means of informing and raising awareness of the consequences of individual actions (e.g. selection of ornamental garden species) when living beside

or close to natural areas. Awareness of the potential impact of introduced non-native species in natural-urban interfaces can promote a greater demand for native species, which also has

multiple advantages for both the user and the environment86. Scientists have an important role to play in achieving this goal; for example, by leading citizen science projects and promptly

communicating their research results to the public, thus constructing a two-way process that should be strengthened over time. This process could be especially important in the case of

incipient plant invasions since people can receive a timely warning about the potential risks of invasive species before they are perceived as a natural component of the landscape and become

valued. METHODS STUDY AREA The abrupt longitudinal precipitation gradient, moisture availability and temperature of Andean Patagonian brings about a transition in vegetation from humid

forests in the west to steppe environments in the east87. The study area is covered mainly by plants of the Subantarctic biogeographic province and, to a lesser extent, the High Andean and

Patagonian biogeographic provinces88. The Subantarctic province is characterized by temperate and cold forests, both deciduous and evergreen, especially conifers and southern beeches of the

genus _Nothofagus_; the High Andean province is characterized by a dominance of xerophytic grasses and creeping or cushion dicotyledons; and the Patagonian province is represented by

ingressions of the Patagonian steppe with scattered low compact shrubs and abundant bare soil – the grasses found here are mainly low88. In Andean Patagonia there are at least 51 PAs (Table

1) with different jurisdictions, zoning, and degrees of protection. Most PAs are intermingled in a mosaic with different types of land use. For example, one of the largest PAs, Parque

Nacional Nahuel Huapi (710.000 ha), is spread over several municipalities whose urban fabric is in close contact with areas of natural vegetation. The majority of these municipalities are

tourist areas (e.g. San Carlos de Bariloche, Villa La Angostura, San Martín de Los Andes), which leads to high connectivity with other urbanizations, increasing the likelihood of spreading

introduced species and, therefore, generating incipient invasions. STUDY SPECIES _Juniperus communis_ L. (common juniper, enebro; Cupressaceae) grows as a shrub or upright tree (up to 12 m

high) but can also acquire a prostrate form, presumably in response to environmental conditions57. The species is usually dioecious and reproduces predominantly by sexual means89. Rooting of

decumbent branches occurs in areas with an oceanic climate although it is not clear whether these branches survive when the original shrub dies57. Female individuals produce axillary green

globose strobiles, which turn bluish-black when mature57. Cones present unusually fleshy and fused scales that give it a berry-like appearance and take two to three years to mature66 (Fig. 

1). Therefore, reproductive female plants can carry fruits at different stages of maturity all year round57. The native range of this species is Panarctic, occurring from the southern Arctic

to about 30° latitude in North America, Europe, and Asia90. In terms of climate, _J. communis_ occupies very different environments, with limitations due to cold (Arctic and Polar and

Northern Urals), drought (Mediterranean and Southern Urals), or high soil moisture (Eastern Alps)90. Its growth rate is strongly controlled by temperature and limited by soil moisture90.

Moreover, high temperatures can decrease its seed viability, particularly by disrupting the growth of the pollen tube and female gametophyte, as well as fertilization91. Since ancient times

this species has been widely used for culinary, medicinal, and ornamental purposes92,93. In Andean Patagonia, local people harvest the fruits from natural populations and sell them to gin

production companies located in the region and other parts of the country. There is a record of the species entering this region in the 1930s, when it was introduced along with other

non-native conifers to be cultivated for ornamental purposes on Isla Victoria (northwestern Andean Patagonia)55. SAMPLING DESIGN—_JUNIPERUS COMMUNIS_ IN ANDEAN PATAGONIA To describe the

invasion of _J. communis_ we compiled records of its location (latitude and longitude) in PAs and their interface areas in the Andean Patagonian region (Argentina). Data were obtained

through field surveys, a literature review, and the contributions of citizens. For each data source, a set of additional variables to describe the invasion and the environment were also

recorded. The number and type of variables depended on the data source. In autumn 2022 we carried out field surveys in an area that encompassed protected areas and their urban-natural

interface areas (− 40.63, − 42.97; − 71.87, − 71.65). As we traveled along main and secondary roads (paved and gravel roads, respectively) and walking trails, we searched for _J. communis_

individuals (sampling point). At each sampling point (with at least 1 km between points) we registered: species location, habitat type (steppe, shrubland, forest, or other), environment type

(natural, rural, or urban), the abundance of individuals (single: 1, low: 2–10, medium: 11–100, and high: > 100), the spatial configuration of the individuals (thicket, isolated, or

both), the main woody species, whether the species was occurring naturally or not (e.g. ornamental), and if there were individuals with fruits and seedlings (< 0.25 m58). The presence of

fruits and seedlings was considered a proxy for reproductive potential. In addition, to compile a set of data we reviewed the scientific literature and literature specializing in the

regional flora (books and technical reports) that reported the occurrence (location) of _J. communis_. In May 2022 we searched the scientific literature on Scopus using the following terms:

“Juniperus AND communis OR enebro OR juniper AND Patagonia AND Argentina”. The reference lists from the articles found were also searched for other relevant publications not found in the

initial search. In addition, to know when and how many times the species has been registered as naturally occurring in the region, and how it was recognized in terms of invasive status, we

looked for articles that reported _J. communis_ as part of the natural vegetation; we registered: the publication year, whether the species was the focus of the article (i.e. if it was

intentionally selected to be studied or not), its recognized status (e.g. introduced, invasive species), the reason why it was included in the study, and if it was registered in a protected

area. During autumn 2022 we also made a call to citizens through social networks, requesting data on the location of _J. communis_, along with information on how to identify it. We enabled a

number of WhatsApp accounts for citizens to send their records to, since this application was one of the preferred ways for people in Argentina to report species sightings23. We asked

citizens to give the species’ location, provide a picture of the plant to verify its identity, and report whether any observed plant had fruits. The fruiting plants could be easily detected

by citizens; it is unlikely they would be confused with the similar fruits of common woody natives (i.e. species with small, rounded, purple fruits) whose fructification period ended in

midsummer, before the survey period64. From both the citizen contributions and the literature review methods we obtained extra information on, for example, species abundance and habitat

type. We incorporated these data into the database and indicated in the results section the number of records for each variable presented. POTENTIAL DISTRIBUTION AND BIOCLIMATIC MATCHING To

estimate the potential distribution of _J. communis_ based on climatic parameters that define the habitat suitability of Andean Patagonia, Argentina, we constructed Species Distribution

Models (SDMs) for each species using the Maxent software94, a maximum entropy modeling method that generates a continuous binomial probability distribution of habitat suitability. For this

we used the records obtained through field sampling and citizen science (Table S2), and also records that we downloaded from the gbif database for _J. communis var. communis_ updated to May

31, 2022. All available worldwide occurrences in gbif were considered; however, only occurrences within its native range were found. In total we obtained 133,981 records95, of which 40,263

were complete with geographic coordinates and were used for habitat suitability modeling. We used the 19 bioclimatic variables available in WORLDCLIM version 2.0 as environmental predictors

for the model96. However, since many of Worldclim’s bioclimatic variables are highly correlated, to avoid errors generated by data multicollinearity, we chose 4 bioclimatic variables:

precipitation of the coldest quarter, precipitation of the warmest quarter, mean temperature of the coldest quarter and mean temperature of the warmest quarter. Our selection criteria for

the variables were based on Pearson's correlation analysis (r < 0.7)97 and on relevant bio-ecological knowledge of the species90,91,98,99. We used layers with a resolution of 2.5 min

for these variables. The model was developed using 75% of the location data, while the remaining 25% was used to validate the model. The algorithm was run with 1000 iterations, through

which MaxEnt increases the model gain by modifying the coefficient of a single feature as a function of the input environmental data. The accuracy of the model was tested using the area

under the curve of the receiver operating characteristic (ROC)100. The contribution of each variable to the final model was determined by randomly permuting the values of that variable

between training points (both presence and background) and measuring the resulting decrease in the area under the curve. Values were normalized to obtain percentages. The relative strength

of each predictor variable was assessed using the Maxent Jackknife test of variable importance. This test shows the importance of the environmental variables by detecting (i) the variable

with the greatest explanatory power, and (ii) the variable with the greatest amount of unique information (not contained in the other variables). Finally, to test whether the observed

bioclimatic characteristics coincide between the PAs of the introduced area in Andean Patagonia (200 randomly selected sites) and records from the native range, we quantitatively compared

the values of the 4 environmental variables included in the model. Values for the native range and the PAs of Andean Patagonia were extracted with the point sampling tool in Qgis and

compared using the Anderson–Darling test. The QGIS version 2.18 spatial analysis software was applied to edit and process all the maps shown in this work. All the reported models, tests, and

graphs were performed in R101. ETHICAL APPROVAL The use of plant parts in the study complies with international, national, and institutional guidelines. DATA AVAILABILITY The data used for

the SDM, collected in the framework of this work, are available in the supplementary material (Table S2). All other data generated and/or analyzed during this study are available from the

corresponding author on reasonable request. CHANGE HISTORY * _ 13 JULY 2023 A Correction to this paper has been published: https://doi.org/10.1038/s41598-023-38496-w _ REFERENCES * Foxcroft,

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INFORMATION AUTHORS AND AFFILIATIONS * Investigaciones de Ecología en Ambientes Antropizados, Instituto de Investigaciones en Biodiversidad y Medioambiente (CONICET-UNCo), R8400, S. C.

Bariloche, Argentina Jorgelina Franzese * Grupo de Genética Ecológica, Instituto de Investigaciones en Biodiversidad y Medioambiente (CONICET-UNCo), Evolutiva y de la Conservación, R8400, S.

C. Bariloche, Argentina Ramiro Rubén Ripa * Instituto de Evolución, Ecología Histórica y Ambiente (CONICET-UTN), San Rafael, Mendoza, Argentina Ramiro Rubén Ripa Authors * Jorgelina

Franzese View author publications You can also search for this author inPubMed Google Scholar * Ramiro Rubén Ripa View author publications You can also search for this author inPubMed Google

Scholar CONTRIBUTIONS R.R. and J.F. conceived and designed the study and conducted the methodology. R.R. performed statistical analysis and figures. J.F., with contributions from R.R.,

wrote the main manuscript text. CORRESPONDING AUTHOR Correspondence to Jorgelina Franzese. ETHICS DECLARATIONS COMPETING INTERESTS The authors declare no competing interests. ADDITIONAL

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visit http://creativecommons.org/licenses/by/4.0/. Reprints and permissions ABOUT THIS ARTICLE CITE THIS ARTICLE Franzese, J., Ripa, R.R. Common juniper, an overlooked conifer with high

invasion potential in protected areas of Patagonia. _Sci Rep_ 13, 9818 (2023). https://doi.org/10.1038/s41598-023-37023-1 Download citation * Received: 13 March 2023 * Accepted: 14 June 2023

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