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Article

Cataloging the Alien Flora of the Costa Brava (NE Iberian Peninsula): New Findings and Updates in a Plant Invasion Hotspot

1
Botanical Institute of Barcelona (IBB), CSIC-CMCNB, Passeig del Migdia s/n, 08038 Barcelona, Spain
2
Escuela de Ciencias Ambientales, Universidad Espíritu Santo (UEES), Samborondón 091650, Ecuador
*
Authors to whom correspondence should be addressed.
Diversity 2025, 17(3), 160; https://doi.org/10.3390/d17030160
Submission received: 3 January 2025 / Revised: 20 February 2025 / Accepted: 21 February 2025 / Published: 25 February 2025
(This article belongs to the Section Plant Diversity)

Abstract

:
The Costa Brava, a Mediterranean coastal strip spanning ca. 250 km (in NE Spain), is a hotspot of plant invasions. To address this issue, the LIFE medCLIFFS project was launched to improve the management of invasive species along sea cliffs in the region. The initial phase of the project involved designing more than 100 transects distributed across the coastline, each 1 km in length, which were to be monitored by volunteers. During the fieldwork for transect design, we identified a total of 126 alien species. This is a substantial number, considering that the observations were confined to the transect areas. Notably, among the 126 observed taxa, 24 represented floristic novelties: 7 were new to the Costa Brava, 7 to the province of Girona, 4 to Catalonia, 1 to Spain, 1 to the Iberian Peninsula, and 4 to Europe. Additionally, we provide information on other alien taxa that, although not novel for the study area, could be significant for control and management efforts. These observations provide a foundation for compiling a comprehensive catalog of the alien flora of the Costa Brava.

1. Introduction

Biological invasions are one of the main causes of biodiversity loss on the planet. Non-native invasive species have contributed, solely or together with other drivers, to 60% of documented global animal and plant extinctions [1]. Regrettably, biological invasions, far from being contained, seem to be worsening, with recent studies showing an increasing trend in both the number of first records [2,3] and damage/associated costs [4,5]. The introduction of new alien plant species can disrupt local natural systems and seriously compromise the viability of native species [6,7,8]. In recent decades, the main pathway for these plants has been through their importation for use in gardening, whether in public or private gardens [9,10]. This is especially evident in areas with a high human population density, particularly in coastal areas, such as the Costa Brava of NE Spain. In such an area, the intensity of propagule pressure—in terms of magnitude and affected area [11,12,13]—plays a key role in the invasion success, particularly in the case of certain species.
Likely as a result of the interaction of several factors—including complex topography, deep habitat transformation and extensive touristic development—the Costa Brava has become a hotspot of plant invasions, arguably the most significant in the whole Iberian Peninsula [14]. Despite the severity of the threats posed by biological invasions in the region, critical knowledge gaps remain. These include limited information on the species involved, the extent of the areas invaded, or the magnitude of their ecological impacts.
To prevent further introductions of alien plant species and enhance the management of existing invasive species, our research institution, the Botanical Institute of Barcelona, is leading a project within the European Union’s LIFE Programme. This project focuses on the Habitat of Community Interest (HCI) 1240—Mediterranean sea cliffs and rocky shores hosting endemic Limonium species. The initial phase of this project, LIFE medCLIFFS [15], involved the design of over 100 transects of ca. 1 km each distributed along the entire Costa Brava coastline. Since early 2023, volunteers from the LIFE medCLIFFS Volunteers’ Network (LmVN) have been conducting annual surveys of these transects to detect the presence of invasive plants and monitor the spread of previously detected species [16].
During the design of the transects in early 2022, we discovered various alien species novelties, found along the volunteer monitoring paths or nearby areas, primarily subspontaneous occasional species often originating from garden escapes or the dumping of gardening waste. By the project’s conclusion in 2026, we aim to compile a comprehensive list of alien species in the Costa Brava, largely based on observations that cover a broader area than the transects monitored by the LmVN.
Given the number of new species documented across the Costa Brava, we present a brief overview of the location and geographical significance of these species. Therefore, the present work aims to (1) provide detailed information on the new findings (Section 3.2), including new species for the studied area (Costa Brava), the province encompassing it (Girona), Catalonia, Spain, the Iberian Peninsula, and Europe; and to (2) add some considerations about certain neophytes already known in the Costa Brava (Section 3.3), either due to their invasive behavior in the study area—some causing severe damage to coastal ecosystems—or because, while not considered invasive, they deserve special attention as they have been observed only sporadically and could easily be overlooked.

2. Materials and Methods

2.1. Study Area

The Costa Brava is a coastal strip extending from the French–Spanish border to Blanes, a small city located about 60 km northeast of Barcelona. This region, spanning approximately 250 km of coastline, is well-known for its rugged appearance, characterized by numerous sea cliffs (brava in Catalan means “rugged” or “wild”), which contrast sharply with the nearby by flat coastlines with long, sandy beaches.
The study area comprised 106 transects defined along the Costa Brava coastline (Figure 1), primarily designed for annual monitoring of 33 pre-defined invasive alien plant species by volunteer participants of the LIFE medCLIFFS project [16]. Each transect is approximately 1 km long, resulting in a total of about 106 km of monitorized coastline. These transects follow coastal trails that are predominantly, but not exclusively, located in the HCI 1240, the target habitat of this LIFE project. Additionally, some plant observations were performed in areas immediately adjacent to the transects.

2.2. Data Gathering and Plant Identification

All observed taxa were either vouchered (particularly for the most accessible locations or those plants with not many difficulties to press) or extensively photographed (additional pictures are available upon request) to facilitate taxonomic identification. iNaturalist (https://www.inaturalist.org/) records were also created for at least one population of each species. Identification was based on the authors’ expertise, relevant floras, specialized monographs (e.g., Agaves of continental North America [17]), recent phylogenetic articles, and consultation with experts in certain taxa (e.g., Gideon F. Smith from Nelson Mandela University for Aloe). In addition, to ease identification we also observed the living specimens of several taxa included in the present work that are cultivated in the Botanical Garden of Barcelona (e.g., Agave parryi, Asparagus falcatus, Echium candicans, Furcraea selloana, Heptapleurum arboriccola, Myoporum laetum). The names of taxa have been updated according to Tropicos [18] and Plants of the World Online (POWO) [19]. The authorities of plant names follow the International Plant Names Index [20]. The names of families are based on the APG IV [21], with subfamilies indicated when relevant.
The locations of our observations are provided as UTM (Universal Transverse Mercator) coordinates, with a precision level of 100 × 100 m (ETRS89 datum). As all observations are located within the Spanish province of Girona, the code of the UTM zone (31T) is not indicated.
The identification of the species is based on published literature and reliable sources. When relevant, we provide these references, as specified in the bibliography. While citizen science platforms offer valuable observational data, we acknowledge that such records may require further verification to ensure accuracy (e.g., correct identification, wild vs. cultivated status). Therefore, in this study, citizen science contributions were considered Supplementary rather than definitive sources.
For all identified species, we provide the following information. First, the native range is determined based on the biogeographical world zonation described by Wallace [22] and Takhtajan [23], and includes the following categories: Australia and New Zealand, Cosmopolitan, Holarctic, Nearctic, Nearctic tropical (i.e., all Americas), Neotropical, South America, New Zealand, Palearctic, Palearctic tropical (i.e., the Old World), Paleotropical, Paleotropical and South Africa, Pantropical, South Africa, and artificial origin. Second, the plant growth type is classified into one of ten categories: annual grasses, annual forbs, perennial grasses, perennial forbs, bulbous monocots, aquatic plants, climbers, succulent plants, shrubs and trees. Third, the introduction pathway is assigned to one of six categories: agriculture, gardening, medicinal, soil stabilization, textile, and trade contaminant. Fourth, the intentionality of the introduction is specified as either deliberate or unintentional. Fifth, the invasion status is assessed for both the Costa Brava and the entirety of Catalonia, using three categories—casual, naturalized, or invasive—based on the definitions and criteria provided in [24]. Finally, the type of floristic novelty is described for each taxon, with species classified as newly reported for one of the following areas: Costa Brava (but present in surrounding areas), the province of Girona, Catalonia, Spain, the Iberian Peninsula, or Europe.

3. Results and Discussion

3.1. Alien Species Richness Detected During the Transect Design: Regional Significance

We observed a substantial number of alien plant taxa (up to 126; Supplementary Table S1) when designing the transects for the LIFE medCLIFFS project. Whilst most have been already recorded in the Costa Brava, and some are relatively common plants, we detected up to 24 floristic novelties within these few square kilometers. This number of alien plant taxa should be regarded as very high for two main reasons: (1) compiling this list of plants was not the primary objective of the fieldwork, which focused on designing the aforementioned transects; therefore, we did not carry out neither an intensive nor structured survey; and (2) the transects, although representative of the coastal cliffs of the Costa Brava, excluded many other habitats types (e.g., large sandy beaches, wetlands, agricultural areas, close forests or roadsides), and account for less than 2 km2 (assuming that a transect is generally 10–15 m wide). When comparing our study area to the smallest region encompassing the Costa Brava with available data on alien species (coastal Catalonia, with 933 taxa in 14,000 km2 [25]), it becomes evident that the cliffs of Costa Brava can be regarded as a hotspot for alien plants.
The 24 floristic novelties documented in this study represent a noticeable increase in our knowledge of the regional alien flora. These novelties include seven new species for the Costa Brava, another seven for the province of Girona, four for Catalonia, one for Spain, one for the Iberian Peninsula, and four for Europe (details provided below). Most of these floristic novelties were observed in the transects located in the southern half of the Costa Brava (Figure 1). This fact aligns with expectations, as this area hosts the highest population density in the region, and human population density is a strong predictor of alien species richness [26].
Although most of these newly detected plant taxa are still occurring as casual (20 cases out of 24), some are already established in the Costa Brava and soon could become very problematic, as seen elsewhere (e.g., Cylindropuntia imbricata). Half of the identified floristic novelties are of American origin (12 out of 24; Table 1), a proportion notably higher than the reported alien flora of coastal Catalonia (32.6% [25]). This discrepancy is likely due to the fact that most of the 24 taxa were introduced for ornamental purposes (23 out of 24 cases, with Nicotiana tabacum being the sole exception, introduced for agricultural purposes). American plants are particularly common in the ornamental flora of Spain [27], which can be attributed to Spain’s historical role as the former metropole of most of colonial America.
It is not surprising to find such a wide array of taxa used for gardening, as most of the cultivated plants within the domestic gardens of the Costa Brava are alien [28]. As expected for plants mostly escaped from nearby gardens (the urbanized areas of the Costa Brava are characterized by high concentrations of secondary single-family houses with domestic gardens [29]), there are very few short-lived herbs among the novelties. In contrast, nearly half (11 of 24; Table 1) are succulent taxa, which can be related to the modern uses of these plants in the Mediterranean coastal areas (i.e., xerogardening [25]).
Table 1. Attributes for the 24 new taxa detected in our study area in Costa Brava. (1) “Cat.”: taxa that appear in the catalog of plants of Catalonia are indicated with a plus sign [30]; (2) novelty (“Nov.”): first observed in CB (Costa Brava), GI (province of Girona), CA (Catalonia), SP (Spain), IP (Iberian Peninsula) or EU (Europe); (3) native range: ART (Artificial), AUS (Australia), AUS-NZ (Australia and New Zealand), NAR (Nearctic), NAR-TRO (Nearctic tropical), NTRO (Neotropical), NZ (New Zealand), PLAR (Palearctic), PLAR-TRO (Palearctic tropical), PLTRO (Paleotropical), PLTRO-SAF (Paleotropical and South Africa), PNTRO (Pantropical), SAF (South Africa), and SAM (South America); (4) family; (5) functional traits (“Fun. traits”) based on Aymerich & Sáez (2019): AFs (annual forbs), BU (bulbous monocots), CLs (climbers), PFs (perennial forbs), PGs (perennial grasses), SHs (shrubs), SU (succulent plants), and TRs (trees); (6) introductory pathway (“Intr. Path.”): A (agriculture), G (gardening) and M (medicinal); (7) intentionality introduction (“Int. intr.”): D (subspontaneous, deliberate) and A (adventive, unintentional); (8) invasion status (“Inv. status”) in Catalonia (and in the Costa Brava in parentheses): C (casual), N (naturalized), and I (invasive). Invasion status in Catalonia is taken from Sáez & Aymerich [30] if present in their catalog.
Table 1. Attributes for the 24 new taxa detected in our study area in Costa Brava. (1) “Cat.”: taxa that appear in the catalog of plants of Catalonia are indicated with a plus sign [30]; (2) novelty (“Nov.”): first observed in CB (Costa Brava), GI (province of Girona), CA (Catalonia), SP (Spain), IP (Iberian Peninsula) or EU (Europe); (3) native range: ART (Artificial), AUS (Australia), AUS-NZ (Australia and New Zealand), NAR (Nearctic), NAR-TRO (Nearctic tropical), NTRO (Neotropical), NZ (New Zealand), PLAR (Palearctic), PLAR-TRO (Palearctic tropical), PLTRO (Paleotropical), PLTRO-SAF (Paleotropical and South Africa), PNTRO (Pantropical), SAF (South Africa), and SAM (South America); (4) family; (5) functional traits (“Fun. traits”) based on Aymerich & Sáez (2019): AFs (annual forbs), BU (bulbous monocots), CLs (climbers), PFs (perennial forbs), PGs (perennial grasses), SHs (shrubs), SU (succulent plants), and TRs (trees); (6) introductory pathway (“Intr. Path.”): A (agriculture), G (gardening) and M (medicinal); (7) intentionality introduction (“Int. intr.”): D (subspontaneous, deliberate) and A (adventive, unintentional); (8) invasion status (“Inv. status”) in Catalonia (and in the Costa Brava in parentheses): C (casual), N (naturalized), and I (invasive). Invasion status in Catalonia is taken from Sáez & Aymerich [30] if present in their catalog.
TaxonCat.Nov.Native RangeFamilyFun. TraitsIntr. Path.Int. Intr.Inv. Status
Agave lurida+GINTROAsparagaceae (Agavoideae)SUGDC (C)
Agave parryi EUNAR-TROAsparagaceae (Agavoideae)SUGDC (C)
Aloe glauca × A. speciosa EUSAFAsphodelaceaeSUGDC (C)
Aloe striata CAPLTRO-SAFAsphodelaceaeSUGDC (C)
Aloe vera+CBPLARAsphodelaceaeSUG & MDC (C)
Asparagus aethiopicus+GISAFAsparagaceaeCLGDC (C)
Asparagus falcatus IPSAFAsparagaceaeSHGDC (C)
Baculellum articulatum CASAFAsteraceaeSUGDC (C)
Bougainvillea glabra+GINTRONyctaginaceaeCLGDC (C)
Brugmansia aurea SPSAMSolanaceaeSHGDC (C)
Cocculus laurifolius EUPLAR-TROMenispermaceaeTRGDC(C)
Cylindropuntia imbricata+CBNAR-TROCactaceaeSUGDN (N)
Echinopsis oxygona+CBSAMCactaceaeSUGDC (C)
Echium candicans+CBPLARBoraginaceaePFGDC (C)
Ficus pumila CAPLAR-TROMoraceaeCLGDC (C)
Furcraea selloana GINTROAsparagaceae (Agavoideae)SUGDC (C)
Heptapleurum arboricola EUPLAR-TROAraliaceaeSHGDC (C)
Myoporum laetum+GIAUS-NZScrophulariaceaeTRGDN (N)
Nicotiana tabacum+CBNTROSolanaceaeAFADC (C)
Opuntia canterae+CBNTROCactaceaeSUGDN (N)
Phormium tenax CANZAsphodelaceae (Hemerocallidoideae)PFGDC (C)
Selenicereus undatus+CBNTROCactaceaeSUGDC (C)
Washingtonia filifera+GINTROArecaceaeTRGDC (N)
Wigandia urens+GINTRONamaceaeSHGDC (C)
The research presented in this paper also highlights the importance of conducting fieldwork, which is essential for the early detection of invasive alien species and the implementation of the Early Detection and Rapid Response (EDRR) strategy [31].

3.2. New Species Reported in the Studied Area

Agave lurida Aiton [incl. Agave vera-cruz Mill.] (Asparagaceae subfam. Agavoideae)
Lloret de Mar, near the residential area called “La Tortuga”, DG888164, 76 m, a single adult plant and a juvenile in an open area of a pinewood, 24 February 2022, J. López-Pujol, C. Gómez-Bellver, J. Martínez-Fuentes & N. Nualart, vidi vivam, https://www.inaturalist.org/observations/107446643 (accessed on 19 February 2025). Santa Maria de Llorell, DG923175, 10 m, a few adult plants surrounded by Smilax aspera, 24 February 2022, J. López-Pujol, C. Gómez-Bellver, J. Martínez-Fuentes & N. Nualart, vidi vivam (Figure 2A).
New for the province of Girona. An elegant agave endemic to Oaxaca in Mexico [17], although there it could be extinct in the wild [32]. Previously reported in a few localities in the southern half of the coastal area of Catalonia (Salou [33]; Sitges: BC-956093/BC-956094, J. López-Pujol & N. Nualart, 4 January 2016; Roquetes: BC-973174, C. Burguera, 30 August 2018; Barcelona (Montjuïc): vidi vivam, J. López-Pujol & N. Nualart, 3 September 2020).
Agave parryi Engelm. (Asparagaceae subfam. Agavoideae)
Sant Feliu de Guíxols, el Fortim, east side of the port, EG029253, 9 m, an individual located on a steep slope surrounded by other subspontaneous plants such as Agave americana, Cylindropuntia imbricata and Crassula ovata, 11 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, BC-FV-983955, https://www.inaturalist.org/observations/262232315 (accessed on 19 February 2025) (Figure 2B).
New for Europe. A small agave native to the southwestern United States and northwestern Mexico [17], with different varieties and forms. It also resembles Agave flexispina Trel., from northeastern Mexico, with larger and stronger teeth and spines. Our plant probably corresponds to the variety huachucensis (Baker) Little [34] (see image in POWO [35]). It is sold as an ornamental plant in Catalonia (C. Gómez-Bellver & J. López-Pujol, pers. obs.), but we have rarely seen it growing in gardens. We are not aware of this species being subspontaneous in the rest of Europe [36,37].
Aloe glauca Mill. × A. speciosa Baker (Asphodelaceae)
Sant Feliu de Guíxols, at the viewpoint of les Modistes, EG023244, 25 m, a few plants were observed growing alongside Opuntia ficus-indica, near the coastal area, 11 March 2022, C. Gómez-Bellver & J. López-Pujol, vidi vivam, https://www.inaturalist.org/observations/262144176 (accessed on 19 February 2025). Towards the south in Ses Ulleres, EG013246, 69 m, some scattered plants in a slope facing the sea coast, 25 October 2022, C. Gómez-Bellver & N. Ibáñez, vidi vivam (Figure 2C).
New for Europe. This hybrid involves two South African species with disjunct areas: Aloe glauca Mill. is found in arid areas on Western and Northern Cape provinces, while A. speciosa Baker is primarily distributed in the Eastern Cape province. However, their ranges come very close in the Swellendam region [38]. The recent CITES checklist for worldwide Aloe spp. does not consider the presence of this hybrid either in the wild or in nurseries. The identity of this plant was confirmed by Gideon Smith (pers. comm.), who mentioned occasional spontaneous hybridization between A. lineata and A. speciosa. Notably, A. lineata features distinct longitudinal stripes, a characteristic partially inherited by the hybrid. However, the plants discovered in Costa Brava either lack visible stripes or have them only faintly.
A recent review on the presence of Aloe × caesia Salm-Dyck [A. arborescens Mill. × A. ferox Mill.] in Catalonia [39] reported some subspontaneous plants in Pau (Alt Empordà county in Girona province). They were attributed to A. × caesia based on their small clusters of leaf rosettes, contrasting with the solitary rosette of A. ferox. However, the long, narrow, unbranched inflorescences illustrated in [39] (p. 332) differ significantly from the description of the genuine Aloe × caesia [40]. This morphology aligns more closely with A. glauca × A. speciosa.
Aloe striata Haw. (Asphodelaceae)
Lloret de Mar, above Freu d’en Sitre, DG885164, 27 m, two plants, one in bloom, on a slope below the path, 24 February 2022, C. Gómez-Bellver, J. López-Pujol, J. Martínez-Fuentes & N. Nualart, vidi vivam, https://www.inaturalist.org/observations/262145544 (accessed on 19 February 2025) (Figure 2D).
New for Catalonia, likely the second report for the Iberian Peninsula. This succulent plant, native to South Africa [38], is not commonly cultivated in gardens. This species was previously observed in Valencia province [41], and it is regarded as casual in Italy [42].
Aloe vera (L.) Burm. f. (Asphodelaceae)
S’Agaró, near Cala de Vaques, EG049269, 7 m, a single plant in a rocky place among a population of Carpobrotus sp., 11 March 2022, J. López-Pujol, vidi vivam, https://www.inaturalist.org/observations/108372455 (accessed on 19 February 2025) (Figure 2E).
New for the Costa Brava, where its presence was not previously recorded [43,44]. Native to the Socotra Island, east of the Horn of Africa. Very popular for its medicinal uses, even in Europe.
Asparagus aethiopicus L. (Asparagaceae)
Blanes, Cala Sant Francesc, DG841142, 16 m, 18 February 2022, C. Gómez-Bellver, J. López-Pujol, N. Ibáñez & N. Nualart, vidi vivam, https://www.inaturalist.org/observations/262300966 (accessed on 19 February 2025) (Figure 2G). Lloret de Mar, Caleta d’en Trons. DG888164, 12 m, 24 February 2022, C. Gómez-Bellver, J. López-Pujol, J. Martínez-Fuentes & N. Nualart, vidi vivam. Roses, Cala de Canyelles Petites, EG160772, 6 m, 28 February 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, vidi vivam. Sant Feliu de Guíxols, Puig de les Forques, EG033252, 15 m, 11 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Nualart, vidi vivam. Castell Platja d’Aro, S’Agaró, Cala de Vaques, EG049269, 11 m, 11 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Nualart, vidi vivam. In all these localities, only one or a few individuals were observed, having probably escaped from nearby gardens.
New for the province of Girona. This South African species was previously observed as subspontaneous in some other localities in Catalonia (Barcelona and Tarragona provinces, e.g., [45], under Asparagus densiflorus (Kunth) Jessop var. sprengeri; C. Gómez-Bellver, pers. obs.) but not in the Costa Brava so far. This species is frequently cultivated in public spaces and private gardens.
Asparagus falcatus L. (Asparagaceae)
Palamós, north of Cap Gros, EG119334, 36 m, a few adult individuals 2–4 m tall, along with some other smaller ones, observed on the margin of a path crossing a pine forest near residential estates, 28 March 2022, C. Gómez-Bellver, J. López-Pujol, N. Besolí & N. Ibáñez, vidi vivam, https://www.inaturalist.org/observations/109741140 (accessed on 19 February 2025) (Figure 2H).
New for the Iberian Peninsula, second report for southwestern Europe. A large thorny South African shrub [46], commonly used as a hedge or for covering garden walls and pergolas. In Europe, this species has only been reported as a casual subspontaneous plant in Italy [36,42].
Baculellum articulatum (L. f.) L.V. Ozerova & A.C. Timonin [Cacalia articulata L. f.; Kleinia articulata (L. f.) Haw.; Senecio articulatus (L. f.) Sch. Bip.] (Asteraceae)
Sant Feliu de Guíxols, el Fortim, east side of the port, EG029253, 10 m, a few groups of plants observed at the base of two large agave specimens (Agave americana f. ingens), located at the edge of a small slope, in fruiting condition, 11 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, BC-983954, https://www.inaturalist.org/observations/108360154 (accessed on 19 February 2025) (Figure 2F).
New for Catalonia. A South African succulent plant cultivated for ornamental purposes. In Europe, it has been observed as being subspontaneous only in a few localities within the Valencian Community [47,48,49,50]. The taxonomic assignment of this species has been notably complex and variable in recent years. Following the conclusions of Ozerova et al. [51], we adopt its reassignment to the monospecific genus Baculellum, which differs from Curio spp. by its deciduous, pachycaulic stem-succulent growth habit, among other distinguishing traits. This criterion has also been adopted in recent works [52].
Bougainvillea glabra Choisy (Nyctaginaceae)
Roses, Cala de Canyelles Petites, EG158771, 31 m, 28 February 2022, a single small plant observed, rooting and flowering, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, BC-FV-983951, https://www.inaturalist.org/observations/107630815 (accessed on 19 February 2025) (Figure 3A).
New for the province of Girona. Native to tropical and subtropical regions of South America, mainly Brazil, Peru and Argentina. Previously observed as being subspontaneous in Catalonia, but only in Barcelona (Montjuïc: J. López-Pujol & C. Gómez-Bellver, vidi vivam, 16 June 2017, BC-983951, 9 June 2020; Vallcarca [53]).
Brugmansia aurea Lagerh. (Solanaceae)
Calonge, on one side of the Andorra avenue, EG070311, 24 m, a large flowering individual observed in a wide hollow with dense vegetation, dominated mainly by Arundo donax, 15 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, BC 983940, https://www.inaturalist.org/observations/108658441 (accessed on 19 February 2025) (Figure 3B).
New for Spain. This ornamental shrub, native to Ecuador [54,55] and, as for the case of Agave lurida, it has been classified as “extinct in the wild” by the IUCN [56]. The species is characterized by its large bell-shaped flowers and is occasionally cultivated in private gardens. A 2022 consultation in the Global Biodiversity Information Facility [57] revealed only one prior record in Europe, a Brugmansia aurea specimen collected in Benicarló (Castelló) [VAL 236656, R. Senar, 23 March 2017, sub B. pittieri (Saff.) Moldenke]. The collector later confirmed the plant’s true identity as B. versicolor Lagerh. An additional report of B. aurea in Italy [58] was erroneously considered as the second observation in Europe, as it relied on the misidentification of the Benicarló (Castelló) specimen. A recent review of GBIF records revealed several iNaturalist observations of B. aurea in southern Europe, though nearly all correspond to cultivated plants. In the Iberian Peninsula, a potential case of a subspontaneous plant was reported in Lisbon (Portugal) [59], requiring further confirmation on the ground. Thus, our record represents a novelty at least for Spain.
Cocculus laurifolius DC. (Menispermaceae) [Pachygone laurifolia (DC.) L. Lian & Wei Wang]
Blanes, southwards the Pinya de Rosa botanical gardens, in an open area where there was an old garden, likely abandoned for decades, DG844147, 43 m, with some trees persisting from former cultivation along with a few young plants, 18 February 2022, C. Gómez-Bellver, J. López-Pujol, N. Ibáñez & N. Nualart, vidi vivam, https://www.inaturalist.org/observations/107172678 (accessed on 19 February 2025) (Figure 3C).
New for Europe. Species of Asian origin, ranging from the Himalayas to Japan [60,61]. It is cultivated in the Iberian Peninsula [61] as an ornamental tree, particularly in public gardens. While Cocculus laurifolius does not appear to have naturalization capabilities, it can persist as a crop remnant with scarce recruitment. We are not aware of any subspontaneous ocurrences of this species elsewhere in Europe [36,62].
Cylindropuntia imbricata (Haw.) F.M. Knuth (Cactaceae)
South of Roses, in the Racó de la Figuerassa, EG175760, 18 m, one individual close to a coastal slope, 28 February 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, vidi vivam. Sant Feliu de Guíxols, at El Fortim, east side of the port, EG029253, 9 m, one single plant on a steep slope, 11 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, vidi vivam, https://www.inaturalist.org/observations/108359125 (accessed on 19 February 2025) (Figure 3D).
New for the Costa Brava. It has previously been reported in other localities in Catalonia, including the south of the province of Girona [63], but not in the Costa Brava.
Echinopsis oxygona (Link) Zucc. ex Pfeiff. & Otto (Cactaceae)
Cadaqués, towards the south, in the Sa Conca beach, EG228810, 9 m, a single plant on a steep slope, 6 March 2024, J. López-Pujol, C. Gómez-Bellver, & N. Ibáñez, vidi vivam, https://www.inaturalist.org/observations/201470180 (accessed on 19 February 2025) (Figure 3E).
New for the Costa Brava. Previously observed subspontaneously in Catalonia in Pira, in the province of Tarragona [64]. This small South American cactus with pinkish or reddish flowers is very similar to another congeneric species, Echinopsis eyriesii, which has white flowers and was observed in a locality in the interior of the province of Barcelona, in Castellbell i el Vilar [63].
Echium candicans L. f. (Boraginaceae)
Sant Feliu de Guíxols, El Fortim, east side of the port, EG029253, 6 m, a small group of plants at the top of an almost vertical slope, 11 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, vidi vivam, https://www.inaturalist.org/observations/262232729 (accessed on 19 February 2025) (Figure 3F).
New for the Costa Brava. An endemic plant to Madeira, it is cultivated in public parks and botanical gardens for its stricking deep blue inflorescences. First recorded as a garden escape in Montjuïc mountain (Barcelona), in 2010 at the edge of a pine forest in the Joan Brossa gardens (S. Pyke, pers. comm.), and in 2017 near the southern entrance of the Mossèn Costa i Llobera gardens [65]. Additional occurrences have been documented in Galicia (Spain) [66], in the Mediterranean coast of France [67], and as naturalized in Italy [42]. Echium candicans thrives in areas with warm or temperate climates, inhabiting ravines, slopes, cliffs and humid forests.
Ficus pumila L. (Moraceae)
Sant Feliu de Guíxols, Cala de ses Mongetes, EG003241, 32 m, some clonal spots of plants clearly escaped from a house in a cleared pine forest, 11 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, vidi vivam, https://www.inaturalist.org/observations/108425728 (accessed on 19 February 2025) (Figure 3H).
New for Catalonia, and likely for the east of the Iberian Peninsula. There is a specimen collected in Blanes without information about dates and collectors, and it is unclear if the plant was wild or cultivated (BCN 68720). Thus, our observation constitutes the first confirmed report of this species. A woody evergreen liana native to East Asia [68], useful for covering the ground, walls or even tree trunks. It has been recorded as a casual non-native plant in the Azores [69], the Canary Islands [70], and Italy [42].
Furcraea selloana K. Koch (Asparagaceae subfam. Agavoideae)
North of Cadaqués, Turó d’en Morell, EG235832, 40 m, two mid-sized plants and a few juveniles, in a road margin, growing among various species escaping from a nearby private garden, 4 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Nualart, BC-FV-983953, https://www.inaturalist.org/observations/262154059 (accessed on 19 February 2025) (Figure 3G).
New for the province of Girona, second report for Catalonia. A Neotropical succulent plant that resembles an agave, native from Mexico to the north of South America [71]. Previously observed in Catalonia only in Barcelona (J. López-Pujol & N. Nualart, 31 August 2020, vidi vivam).
Heptapleurum arboricola Hayata [Schefflera arboricola (Hayata) Merr.] (Araliaceae)
Castell Platja d’Aro, S’Agaró, EG046267, 25 m, a single plant thriving in a crevice in an urban stone wall, 11 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, vidi vivam, https://www.inaturalist.org/observations/108372557 (accessed on 19 February 2025) (Figure 4A).
New for Europe. An evergreen and somewhat climbing shrub native to Taiwan Island [72], very popular within its genus (Schefflera). Recent taxonomic revisions have reassigned this species to the genus Heptapleurum Gaertn. [73]. To our knowledge, this is the first subspontaneous record in Europe [36,74].
Myoporum laetum G. Forst. (Scrophulariaceae)
To the north of Platja d’Aro, in Cala de Calanca, EG069305, 3 m, several subspontaneous individuals on a slope by the sea, probably originating from other cultivated trees, all in an open area, 15 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, vidi vivam, https://www.inaturalist.org/observations/108650753 (accessed on 19 February 2025) (Figure 4B).
New for the province of Girona. A tree native to New Zealand [61], frequently planted in public gardens. Reported as subspontaneous mainly in the coastal areas of Catalonia (Barcelona and Tarragona provinces), the Valencian Community, Murcia and Andalusia (e.g., [30,75]).
Nicotiana tabacum L. (Solanaceae)
Roses, EG150770, 3 m, one well-developed adult in blossom with remnants of fruiting from the previous year, hidden among large coastal rocks, 28 February 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, vidi vivam, https://www.inaturalist.org/observations/107662183 (accessed on 19 February 2025) (Figure 4C).
New for the Costa Brava. This species, though casually found scattered throughout Catalonia [30], has not yet been reported in the Costa Brava. In Catalonia, it was introduced from the Neotropics for tobacco production, and its cultivation has been documented since the late 19th century. However, today its cultivation has either been abandoned or occurs on a very local scale [45].
Opuntia canterae Salm-Dyck (Cactaceae)
Sant Feliu de Guíxols, Puig de les Forques, EG035252, 46 m, a small group of plants on a sea cliff, 11 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, vidi vivam, https://www.inaturalist.org/observations/262152343 (accessed on 19 February 2025) (Figure 4D).
New for the Costa Brava. Previously recorded in Catalonia as Opuntia elata Salm-Dyck [76,77,78,79] but not in the Costa Brava. A recent taxonomic review of the South American Opuntia series Armatae K. Schum. [80] demonstrated that Opuntia elata s.s. is not naturalized in the Mediterranean region or Europe. Instead, species from this group in Catalonia correspond to O. canterae (formerly misidentified as O. elata), O. rioplatensis, and O. monacantha.
Phormium tenax J.R. Forst. & G. Forst. (Asphodelaceae subfam. Hemerocallidoideae)
Blanes, cala Sant Francesc, DG839140, 13 m, a well-developed plant in rocky terrain by the sea, near a detached house where this species is cultivated, 18 February 2022, C. Gómez-Bellver, J. López-Pujol, N. Ibáñez & N. Nualart, vidi vivam, https://www.inaturalist.org/observations/262150201 (accessed on 19 February 2025) (Figure 4F).
New for Catalonia and likely new for the eastern Iberian Peninsula. This species is native to New Zealand [70], where it was traditionally used for fiber. Today, its use as an ornamental plant is widespread worldwide. It is naturalized in the United Kingdom in similar environments [81], and in the northwest of the Iberian Peninsula, in Lugo [82]. Additionally, it also occurs as a subspontaneous species in Italy [42], the Azores, Madeira [69], and the Canary Islands [83].
Selenicereus undatus (Haw.) D.R. Hunt [Cereus undatus Haw.; Hylocereus undatus (Haw.) Britton & Rose] (Cactaceae)
Lloret de Mar, close to Sa Caleta, DG881165, 10 m, a few plants in a rocky cliff with some detached fragments below, 24 February 2022, C. Gómez-Bellver, J. López-Pujol, J. Martínez-Fuentes & N. Nualart, vidi vivam (Figure 4H). Lloret de Mar, Estirador de Fitora, DG888163, 10–30 m, numerous plants cascading down from houses above the path to almost the base of the cliff, 24 February 2022, C. Gómez-Bellver, J. López-Pujol, J. Martínez-Fuentes & N. Nualart, vidi vivam, https://www.inaturalist.org/observations/107447864 (accessed on 19 February 2025). Tossa de Mar, above the Cala Gran, DG943182, 20 m, a single plant at the base of a metallic fence, 24 February 2022, C. Gómez-Bellver, J. López-Pujol, J. Martínez-Fuentes & N. Nualart, vidi vivam.
New for the Costa Brava. Neotropical climbing cactus previously observed in a few other localities in the south of Catalonia [84,85,86].
Washingtonia filifera (Linden ex André) H. Wendl. [incl. Washingtonia robusta H. Wendl.] (Arecaceae)
Blanes, Cala Sant Francesc, DG839141, 6 m, a small individual at the foot of a wall, and Blanes, southwards Cala Sant Francesc, DG839139, 6 m, a well-developed plant ca. 1 m tall on a sea cliff nearby the sea, 18 February 2022, C. Gómez-Bellver, J. López-Pujol & N. Nualart, vidi vivam. North of Lloret de Mar, DG890164, 35 m, a small plant in a crevice, 24 February 2022, C. Gómez-Bellver, J. López-Pujol, J. Martínez-Fuentes & N. Nualart, vidi vivam. Castell Platja d’Aro, S’Agaró, EG003241, 5 m, a single plant ca. 70 cm tall on a rocky slope, and Sant Feliu de Guíxols, near Cala de Ses Mongetes, EG003241, 33 m, several individuals of various sizes, 11 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Nualart, vidi vivam. San Antoni de Calonge, north of Cap Roig beach, EG070306, 5 m, juvenile plants subspontaneous in an artificial slope, and San Antoni de Calonge, Cala del Forn, EG073311, 5 m, a juvenile plant, 15 March 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, vidi vivam. Palafrugell, north of Tamariu, between Les Trones and Sa Negra, EG173468, 8–10 m, two adults ca. 1 m tall, 10–12 m apart, on a sea cliff (Figure 4G), and Begur, Sa Riera beach, EG176406, 6 m, a few subspontaneous juvenile plants near a large palm-tree probably planted, 4 April 2022, C. Gómez-Bellver, J. López-Pujol & N. Ibáñez, vidi vivam, https://www.inaturalist.org/observations/107447864 (accessed on 19 February 2025).
First confirmed record of this species for the province of Girona. A palm tree native to southwestern North America (California, Arizona and Sonora). Recent studies concluded that there is no solid evidence to consider Washingtonia filifera and W. robusta as two different species, as the observed morphological and genetic variation might result from differences in environmental conditions [87,88,89]. While the recent checklist of the vascular plants of Catalonia [30] suggested the presence of this species in the province of Girona (Ruscinic region) in a broad sense, this is the first precise documentation of specific localities. Most of the individuals observed, if not all, likely originated from bird-dispersed seeds [90] from nearby cultivated specimens.
Wigandia urens (Ruiz & Pav.) Kunth (Namaceae)
Blanes, south of the Pinya de Rosa botanical gardens, in an open area, DG845148, 46 m, a small group of plants, 18 February 2022, C. Gómez-Bellver, J. López-Pujol, N. Ibáñez & N. Nualart, vidi vivam, https://www.inaturalist.org/observations/106970273 (accessed on 19 February 2025) (Figure 4E).
New for the province of Girona. A Neotropical garden plant known for its stinging properties upon skin contact. Previously recorded in the coast of Barcelona (Castelldefels: [45]; Montjuïc: C. Gómez-Bellver, N. Ibáñez & P. Farelo, 6 July 2017, BC-990450).

3.3. Considerations About Some Remarkable Neophytes in the Costa Brava

Some species found in the Costa Brava deserve especial attention due to their rarity, taxonomic uncertainties, or because we have observed a highly invasive behavior, causing negative impacts on the biodiversity in coastal habitats and its native plant species.
Cylindropuntia pallida (Rose) F.M. Knuth (Cactaceae)
A densely spiny cactus, invasive in various parts of Catalonia and sporadically found in the southern Costa Brava [43]. We observed a small group of subspontaneous individuals on a rocky wall along the coastal path north of Lloret de Mar (24 February 2022, 26 m. J. López-Pujol, N. Ibáñez & C. Gómez-Bellver, vidi vivam, Figure 5A). Due to its risk to humans and animals, we included this species on the LIFE medCLIFFS invasive species monitoring list. We notified this occurrence to the local authorities and their removal was carried out on 8 July 2022. Additionally, we found a small plant in an open area near the coast in La Montgoda, Lloret de Mar ([91] 23 December 2023, G. Ibáñez, N. Ibáñez & A. Bosch-Guiu) and we also notified its presence to the local authorities.
Delosperma ecklonis (Salm-Dyck) Schwantes (Aizoaceae)
This species is native to the eastern part of South Africa, first reported in Catalonia in Blanes [92]. More recently, we observed this plant in the same locality. In some cases, it is found in areas where Disphyma crassifolium—another South African species more widespread along the coastal region—is also present, with which it can be mistaken due their similar morphology. However, while Delosperma ecklonis has hairy stems and leaves, D. crassifolium is glabrous [93]. The leaves of both species are fleshy and triangular in cross-section, and their flower color ranges from white to pink or deep violet. In its native region, D. ecklonis generally has white or pale pink flowers, whereas D. crassifolium is more variable, though typically pink or violet. As Aymerich [92] already pointed out, this variability can be explained by its ornamental origin.
We observed groups of both species coexisting in rocky areas north of Lloret de Mar, where Disphyma crassifolium exhibited white flowers, while D. ecklonis individuals displayed violet flowers (DG888163, 30 m, 24 February 2022). More recently, we confirmed its presence behind the beach at Cala Jóncols (EG208779, 2 m, 6 March 2023) within the Cap de Creus Natural Park. Additionally, it has been recorded in Tossa de Mar (Figure 5B), where two populations are documented on iNaturalist ([94] 24 June 2023; [95] 5 May 2024, M. Riera).
Drosanthemum candens (Haw.) Schwantes (Aizoaceae)
This South African species was reported by Pyke [96] from sea cliffs at various locations along the Costa Brava (Figure 5C) and it has apparently been confused on several occasions with Drosanthemum floribundum. While there is a general consensus that these are two distinct species, some sources (e.g., POWO) list D. candens as a synonym of D. floribundum. Both species are known as ornamental plants under the common name “queen’s crown”, characterized by their dense, carpet-like growth and numerous showy small flowers, often used for cascading floral displays. However, this commercial name is generally applied to D. floribundum, though it may occasionally refer to D. candens.
According to the literature, in its native area, D. candens is a mat-forming shrublet with grey leaves and cymes of pale pink or white flowers, whilst D. floribundum is a sprawling shrublet with shiny leaves and magenta solitary flowers with a white centre [93,97]. It is also important to note that commercialized plants may differ slightly from their wild counterparts.
Kalanchoe × houghtonii D.B. Ward (Crassulaceae)
This hybrid, allegedly formed by the crossing of Kalanchoe daigremontiana Raym.-Hamet & H. Perrier and K. delagoensis Eckl. & Zeyh., both from Madagascar, has rapidly spread worldwide in just 80 years and is now present on all continents except Antarctica [98]. Most global occurrences correspond to one of its four morphotypes, morphotype A, although morphotype B is also found in some countries; in contrast, morphotypes C and D are restricted to Madagascar (see [99]).
This hybrid was first detected in the Valencian Community about three decades ago [100,101] and has since spread along much of the Iberian Peninsula coastline, including parts of the Costa Brava [102]. We believe that public awareness, including that of environmental authorities, regarding its invasive potential remains low. Nowadays, this nothospecies is clearly expanding, particularly in coastal environments and inland areas with mild climates. Due to its effective clonal propagation (via pseudobulbils and plant fragments), its eradication could be one of the most complicated and expensive among invasive species, as it often forms very dense living carpets (with densities up to 1000–2000 individuals/m2 [98]), which prevent the germination and growth of native herbaceous species.
As a result of our recent fieldwork, we detected 32 localities where this taxon was present, ranging from small groups to populations of hundreds or thousands of individuals (the localities are detailed in the Supplementary Table S2). Competition with the native flora is relevant in some areas (e.g., Roses; Figure 5D). All but one observed locality corresponded to the most common morphotype (A). However, near the s’Arenella lighthouse (in Port de la Selva), we observed morphotype B (Figure 5E), marking the first report of this morphotype (which has smaller leaves than morphotype A along with other differences [99]) in Europe. Approximately one year and half later (May 2023), we located the second European population of this morphotype in Ostia (Italy). Recently, karyological differences have also been observed between the two morphotypes (morphotype A is tetraploid and morphotype B diploid [103]).
Additionally, one of its parental species, Kalanchoe delagoensis, was observed in several locations along the Costa Brava: Blanes, a residential area above Cala de Sant Francesc, DG841143, 58 m, with a few plants in a narrow strip of garden that are apparently subspontaneous, 18 February 2022, C. Gómez-Bellver, J. López-Pujol, N. Ibáñez & N. Nualart, BC-983929; Lloret de Mar, La Tortuga, DG982165, 23 m, with a group of flowering plants in the process of naturalization, 24 February 2022, C. Gómez-Bellver, J. López-Pujol, J. Martínez-Fuentes & N. Nualart, BC-983947; Palamós, Cala de Castell, EG128345, 10 m, with some scattered adult plants in blossom with considerable recruitment, 28 March 2022, C. Gómez-Bellver, J. López-Pujol, N. Besolí & N. Ibáñez, BC-984276.
Opuntia aurantiaca Lindl. (Cactaceae)
We observed this plant towards the south of Tossa de Mar, near the Codolar Tower, DG942181, 22 m, with a few plants close to a metallic fence, 24 February 2022, C. Gómez-Bellver, J. López-Pujol, J. Martínez-Fuentes & N. Nualart, BC-983950, Figure 5F).
This thorny cactus is likely a sterile hybrid capable of clonal propagation. It was previously reported as invasive in the Costa Brava, occurring both near the coastline and in inland areas, including Vilarnadal, Pau and Palafrugell, observed during 2017–2018 [104] and Vilajuïga, where eradication measures have been implemented [105]. It was first recorded in Europe in Caldes de Montbui (Barcelona) [106]. Since then, its rapid expansion has been particularly concerning, with over 15 localities in Catalonia reporting its presence.
Pittosporum tobira (Thunb.) W.T. Aiton (Pittosporaceae)
This shrub or small tree, native to East Asia, was introduced into the Iberian Peninsula in the 19th century as an ornamental plant [27]. In the eastern Iberian Peninsula, it is primarily distributed along the Mediterranean coast, including the Balearic Islands. Japanese pittosporum (pitòspor in Catalan) is frequently cultivated in public gardens and as a hedge. It can be found in disturbed pine forests or near inhabited areas, either as isolated individuals or very scattered, or already naturalized, some of which exhibit invasive behavior, particularly in coastal areas of the Costa Brava.
The effective spread of this species is partly due to the frugivorous birds that consume its fruits and disperse the seeds [107,108]. In Catalonia [109], the Balearic Islands [110] and Valencian Country [111], Pittosporum tobira is recognized as invasive, where several removal efforts have been undertaken. Its high invasive potential often goes unnoticed, as it superficially resembles some native shrubs and trees. For this reason, most people are not aware of its potential ecological impact for the local biodiversity.
We observed that this plant ranges from occasional to invasive in many parts of the Costa Brava, particularly from Blanes to the southern areas of Cap de Creus. In some locations, it forms extensive patches, occupying the majority of the space (Figure 5G).
Senecio angulatus L. f. (Asteraceae)
This South African succulent, creeping and evergreen plant has attractive yellow capitula that forms dense vine tangles and mats. Alongside Kalanchoe × houghtonii, it is one of the most problematic plants in Mediterranean Spain with rapid and aggressive spread. We recorded this species in more than 40 localities along the Costa Brava. Its ability to cover the whole ground and climb over other vegetation poses a severe threat to native species, such as Pistacia lentiscus—llentiscle in Catalan—by outcompeting them for space and resources (see Figure 5H).

4. Conclusions

Prevention, early detection (ED), and rapid response (RR) are the most effective measures to combat invasive species in general [11,30,112]. It is important to (1) understand the ecology and traits of potentially invasive species that could threaten natural habitats and the native biodiversity, (2) avoid their arrival to our territory (prevention), (3) detect them at a very early stage (early detection) if they have already arrived in a given area, and (4) act swiftly (rapid response) before they spread and establish significant populations that are difficult to eradicate. In this context, the information we provide on newly detected species, as well as the details on already known invasive species, should be taken into consideration by the authorities, stakeholders and land managers for informed decision-making.
For decades, numerous alien plant species have been recorded in coastal areas of Mediterranean Spain (e.g., [65,113,114]), with some of them exhibiting significant invasive behavior. This trend is largely attributed to human impact on the environment, including habitat transformation, degradation, and the widespread use of ornamental plants, which often escape from cultivation and establish in natural ecosystems. In this regard, we have particularly observed this phenomenon in areas with high human activity, such as coastal zones and large urban developments.
The Costa Brava has been massively urbanized, largely linked to the expansion of mass tourism since the 1950s. Herein, as a result of a European Union-funded LIFE project, we report several newly documented alien species along this coastal strip. Some represent significant floristic novelties, including new records for Spain (Brugmansia aurea), the Iberian Peninsula (Asparagus falcatus) and even Europe (Agave parryi, Aloe glauca × A. speciosa, Cocculus laurifolius, and Heptapleurum arboricola). Additionally, we provide the first European record of morphotype B of Kalanchoe × houghtonii, along with valuable information on other alien taxa that, while not new for the study area, could aid in their control and management.
This research also underscores the crucial role of fieldwork in the detection of invasive alien species, enabling the implementation of an effective early detection and rapid response (EDRR [31]) strategy. Given that some of the species found as novelties or the neophytes discussed in this study have proven to be invasive elsewhere (e.g., Asparagus aethiopicus [115], Cylindropuntia imbricata [116,117,118], Cylindropuntia pallida [118,119,120,121,122,123] and Kalanchoe × houghtonii [98]), it is essential to continuously monitor this kind of species to enable a rapid response should they exhibit invasive behavior. Additionally, the taxonomic and distributional data presented here, alongside those being gathered through the ongoing 5-year LIFE project, will pave the way for a complete catalog of the alien flora of the Costa Brava. This resource will be essential in helping conservation managers prioritize taxa requiring urgent intervention.
Finally, we would also like to stress that 2 of the 24 species reported here as new to the study area are highly threatened in their native areas. This is not only an excellent example of the “conservation paradox of threatened and invasive species” [124], but illustrates the risk of overharvesting plants for ornamental use (and the associated international plant traffic); a recent study reported that the high demand for wild cacti has led to unsustainable trade, thereby threatening their natural populations [125].

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/d17030160/s1. Table S1: The 126 plant taxa detected in our study area in Costa Brava; Table S2: New localities of Kalanchoe × houghtonii detected in our study area, sorted by data. Locality, UTM, altitude (m), date and authors are provided for each one.

Author Contributions

Conceptualization, C.G.-B., N.N., N.I. and J.L.-P.; methodology, C.G.-B. and J.L.-P.; validation, N.N.; investigation, C.G.-B., N.N., A.B.-G., N.I. and J.L.-P.; data curation, N.N. and N.I.; writing—original draft preparation, C.G.-B. and J.L.-P.; writing—review and editing, C.G.-B., N.N., A.B.-G., N.I. and J.L.-P.; project administration, C.G.-B. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the LIFE20 NAT/ES/001223 project (LIFE medCLIFFS) funded by the European Union.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding author/s.

Acknowledgments

The authors would like to acknowledge Romà Senar for providing information on a Brugmansia aurea voucher specimen, Gideon F. Smith for helping to understand Aloe glabra × speciosa, and M. Riera for his observations of Delosperma ecklonis and G. Ibáñez for her observations of Cylindropuntia pallida both in iNaturalist.

Conflicts of Interest

The authors declare no conflicts of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

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Figure 1. Left: Map of the Costa Brava (outlined in black) showing the distribution of over 1001 km transects (marked in red) designed for volunteer monitoring of alien flora within the LIFE medCLIFFS project. Right: Distribution of the locations of the 24 floristic novelties (marked in green).
Figure 1. Left: Map of the Costa Brava (outlined in black) showing the distribution of over 1001 km transects (marked in red) designed for volunteer monitoring of alien flora within the LIFE medCLIFFS project. Right: Distribution of the locations of the 24 floristic novelties (marked in green).
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Figure 2. (A): Agave lurida, Santa Maria de Llorell. (B): Agave parryi, Sant Feliu de Guíxols. (C): Aloe glabra × A. speciosa, Sant Feliu de Guíxols. (D): Aloe striata, Lloret de Mar. (E): Aloe vera, S’Agaró. (F): Baculellum articulatum, Sant Feliu de Guíxols. (G): Asparagus aethiopicus, Blanes, Cala Sant Francesc. (H): Asparagus falcatus, Palamós. Pictures (A,B,E,F) by Jordi López-Pujol and (C,D,G,H) by C. Gómez-Bellver.
Figure 2. (A): Agave lurida, Santa Maria de Llorell. (B): Agave parryi, Sant Feliu de Guíxols. (C): Aloe glabra × A. speciosa, Sant Feliu de Guíxols. (D): Aloe striata, Lloret de Mar. (E): Aloe vera, S’Agaró. (F): Baculellum articulatum, Sant Feliu de Guíxols. (G): Asparagus aethiopicus, Blanes, Cala Sant Francesc. (H): Asparagus falcatus, Palamós. Pictures (A,B,E,F) by Jordi López-Pujol and (C,D,G,H) by C. Gómez-Bellver.
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Figure 3. (A): Bougainvillea glabra, Roses. (B): Brugmansia aurea, Calonge. (C): Cocculus laurifolius, Blanes. (D): Cylindropuntia imbricata, Roses. (E): Echinopsis oxygona, South of Cadaqués. (F): Echium candicans, Sant Feliu de Guíxols. (G): Furcraea selloana, North of Cadaqués. (H): Ficus pumila, Sant Feliu de Guíxols. Pictures (A,B,E,F) by Jordi López-Pujol and (C,D,G,H) by C. Gómez-Bellver.
Figure 3. (A): Bougainvillea glabra, Roses. (B): Brugmansia aurea, Calonge. (C): Cocculus laurifolius, Blanes. (D): Cylindropuntia imbricata, Roses. (E): Echinopsis oxygona, South of Cadaqués. (F): Echium candicans, Sant Feliu de Guíxols. (G): Furcraea selloana, North of Cadaqués. (H): Ficus pumila, Sant Feliu de Guíxols. Pictures (A,B,E,F) by Jordi López-Pujol and (C,D,G,H) by C. Gómez-Bellver.
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Figure 4. (A): Heptapleurum arboricola, Castell Platja d’Aro. (B): Myoporum laetum, Platja d’Aro. (C): Nicotiana tabacum, Cadaqués. (D): Opuntia canterae, Sant Feliu de Guíxols. (E): Wigandia urens, Blanes. (F): Phormium tenax, Blanes. (G): Washingtonia filifera, Palafrugell. (H): Selenicereus undatus, Lloret de Mar. Pictures (AC,G) by Jordi López-Pujol, (D,F,H) by C. Gómez-Bellver and (E) by N. Ibáñez.
Figure 4. (A): Heptapleurum arboricola, Castell Platja d’Aro. (B): Myoporum laetum, Platja d’Aro. (C): Nicotiana tabacum, Cadaqués. (D): Opuntia canterae, Sant Feliu de Guíxols. (E): Wigandia urens, Blanes. (F): Phormium tenax, Blanes. (G): Washingtonia filifera, Palafrugell. (H): Selenicereus undatus, Lloret de Mar. Pictures (AC,G) by Jordi López-Pujol, (D,F,H) by C. Gómez-Bellver and (E) by N. Ibáñez.
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Figure 5. (A): Cylindropuntia pallida, Lloret de Mar. (B): Delosperma ecklonis, Tossa de Mar. (C): dense carpet of Drosanthemum candens in the Costa Brava. (D): large population of Kalanchoe × houghtonii near Roses. (E): morphotype B of Kalanchoe × houghtonii, Port de la Selva. (F): Opuntia aurantiaca, Tossa de Mar. (G): Pittosporum tobira in sea cliff. (H): dense population of Senecio angulatus. Pictures (A,C,G) and (H) by C. Gómez-Bellver, (B) by M. Riera and (D,E) and (F) by J. López-Pujol.
Figure 5. (A): Cylindropuntia pallida, Lloret de Mar. (B): Delosperma ecklonis, Tossa de Mar. (C): dense carpet of Drosanthemum candens in the Costa Brava. (D): large population of Kalanchoe × houghtonii near Roses. (E): morphotype B of Kalanchoe × houghtonii, Port de la Selva. (F): Opuntia aurantiaca, Tossa de Mar. (G): Pittosporum tobira in sea cliff. (H): dense population of Senecio angulatus. Pictures (A,C,G) and (H) by C. Gómez-Bellver, (B) by M. Riera and (D,E) and (F) by J. López-Pujol.
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MDPI and ACS Style

Gómez-Bellver, C.; Nualart, N.; Bosch-Guiu, A.; Ibáñez, N.; López-Pujol, J. Cataloging the Alien Flora of the Costa Brava (NE Iberian Peninsula): New Findings and Updates in a Plant Invasion Hotspot. Diversity 2025, 17, 160. https://doi.org/10.3390/d17030160

AMA Style

Gómez-Bellver C, Nualart N, Bosch-Guiu A, Ibáñez N, López-Pujol J. Cataloging the Alien Flora of the Costa Brava (NE Iberian Peninsula): New Findings and Updates in a Plant Invasion Hotspot. Diversity. 2025; 17(3):160. https://doi.org/10.3390/d17030160

Chicago/Turabian Style

Gómez-Bellver, Carlos, Neus Nualart, Arnau Bosch-Guiu, Neus Ibáñez, and Jordi López-Pujol. 2025. "Cataloging the Alien Flora of the Costa Brava (NE Iberian Peninsula): New Findings and Updates in a Plant Invasion Hotspot" Diversity 17, no. 3: 160. https://doi.org/10.3390/d17030160

APA Style

Gómez-Bellver, C., Nualart, N., Bosch-Guiu, A., Ibáñez, N., & López-Pujol, J. (2025). Cataloging the Alien Flora of the Costa Brava (NE Iberian Peninsula): New Findings and Updates in a Plant Invasion Hotspot. Diversity, 17(3), 160. https://doi.org/10.3390/d17030160

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