Management strategy based on multi-staff integration target control of lionfish invasion on a South Atlantic oceanic island

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Pereira do Eirado Silva, Lucas Penna Soares Santos, Pedro H. Pereira Cipresso, and 18 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8626827/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 5 You are reading this latest preprint version Abstract The lionfish invasion in the South Atlantic is recent compared to the Caribbean, and long-term control measures are still unknown. In this article, we detail the management actions between 2020 and 2025 in the Fernando de Noronha archipelago, a world heritage site and biodiversity hotspot. A total of 2,660 lionfish individuals were recorded during 58 months of management, noting exponential population growth (345.9% of increase), with the number of records being higher in monitoring (n = 2,258; focused dives) compared to routine (n = 402; recreational dives) operations. Field detection was more related to ecological variables such as deep waters, coral substrate, weak marine currents, and resting fish behavior. High densities were generally observed at 3.9 ind./ha but increased over time and depth gradient, with a maximum value of 60 ind./ha. Individuals averaged 23.17 ± 6.7 cm in total length and 238.2 ± 241.6 g in weight (max.: 460 mm and 1670.0 g), presenting allometric growth (b = 3.22) and high physiological condition (K = 1.43). Lionfish showed high dispersion with 116 sites recorded around the archipelago and well explored areas represented 353.64 ha. Interpolation analysis reached 75,742 (CI 95% 56,202–95,372) individuals regarding local population estimate. Field management proved to be effective locally in maintaining a stable temporal trend with low density averages. This work demonstrates that the use of legal and protocolary adoptions has served efficiently to stabilize and outline lionfish control through standardized actions. Invasive Alien Species Management protocol Monitoring Diving Fernando de Noronha Archipelago. Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Introduction A challenging and growing topic for the conservation of natural environments is the introduction of invasive alien species (i.e. intentionally or unintentionally introduced, as the same, non-native species; hereafter IAS). IAS complies the second leading cause of species extinction in natural environments and the reason for large-scale environmental and socioeconomic changes (Molnar et al. 2008; Bellard et al. 2016; Vilà and Hulme 2018; Giakoumi et al. 2019). Although high-cost, strategies to control IAS presented good results on several cases (Giakoumi et al. 2019), treating as management all the actions needed to prevent, containment, and mitigation (Blackburn et al. 2011). In respect to marine environment, eradication cases are rare, regarding most occasions from local invasions (see Willan et al. 2000; Wotton et al. 2004; Anderson 2005; Genovesi 2005). In established populations, it is more likely to be expected as a good result the suppression directed at decrease densities rates, which can hold the native ecosystem resilience (Green et al. 2014; Usseglio et al. 2017). The connectivity of aquatic environment and larger dynamic conditions, as currents and depth, complicates the full removal IAS, which are dispersed by many anthropogenic vectors (Bax et al. 2003; Johnston et al. 2009; Ojaveer et al. 2015). This includes those organisms that perform active swimming through water motion, overcoming physicochemical barriers by their permeability and invading new places, even distant (e.g. oceanic islands) or on isolated biogeographical provinces (Floeter et al. 2008; Johnston et al. 2009). Management strategies against IAS can proceed in many ways and each approach must consider the interlocution through global and regional protocols, assessing the IAS features to tailor all efforts for successful reduction (Thresher and Kuris 2004; Giakoumi et al. 2019). To note different aspects (e.g. dispersal capacity, movement ecology, distribution in the non-native area) and the level of invasion, managers can adopt different strategies and prioritize resources and technical applications (Hulme 2006). From there, the main groups involved in the IAS problem can adopt well-founded and scientifically supported actions, with the selection of financial and stakeholder networks. For example, lionfish ( Pterois spp.) in the invaded Atlantic areas are usually managed through physical removals with the participation of local divers and fishers (Green et al. 2014; Usseglio et al. 2017). Public participation contributes to the involvement of society, as is the case with recreational divers engaged in citizen-science projects, carrying out biodiversity surveys, monitoring the health of reef environments, and managing IAS (Pocock et al. 2014; Thiel et al. 2014; Branchini et al. 2015; Pasternak et al. 2019; Earp and Liconti 2020; Hermoso et al. 2021). This way of controlling IAS combines standardized protocols with regional planning, which is considered an effective method for broadening management efforts (e.g. Frazer et al. 2012; de León et al. 2013). Native to the Indo-Pacific, the lionfish Pterois volitans and P. miles are well-documented species with their invasion tracked over the last few decades, notoriously through north-southwestern Atlantic Ocean expansion in the USA, through the Caribbean region, and more recently on the Brazilian marine waters (Schofield 2010; Johnston and Purkis 2011; Côté et al. 2013; Ferreira et al. 2015; Luiz Jr. et al. 2013, 2021; Soares et al. 2023; 2025). It is a typical R-strategy case with elevated reproduction rates, a highly dispersive larval stage, and a generalist predator, with wide occurrence in different environments (e.g. reefs, seagrass, estuaries, mesophotic and anthropic reefs) and conditions (e.g. temperature, depth, salinity) (Albins and Hixon 2008; Johnston and Purkis 2011; Claydon et al. 2012; Jud and Layman 2012; Layman and Allgeier 2012; Jud et al. 2015). Population establishment in non-native areas is notably higher, with density rates exceeding native areas several times over (Darling et al. 2011; Kulbicki et al. 2012; Sandel et al. 2015). So, reducing local abundance of lionfish for minimizing the negative impact on non-native areas is the focus of many institutions, and an efficient way of achieving this goal is to involve environmental agencies with the local community in a participatory way. However, most of the data and information on effective management actions and population-reduction measures for lionfish come from the Caribbean, where the invasion has been underway for about 30 years (Claydon et al. 2012; Frazer et al. 2012; Albins and Hixon 2013; de León et al. 2013; Usseglio et al. 2017). Since 2020, several lionfish ( Pterois spp.) sightings across the Southwestern Atlantic coast (~ 4,000km) revealed to have successfully established population, including at least 18 Marine Protected Areas (MPAs) (Soares et al. 2023; 2025). The Chico Mendes Institute of Biodiversity Conservation (ICMBio), a Brazilian government agency and part of the Ministry of the Environment and Climate Change, has as its main function the administration of protected natural areas, such as important MPAs. In addition, the team manages the list of threatened species, implements action plans at national level and deals with the context of IAS and the impacts they cause (ICMBio 2023). Considering the history of invasions in different regions of the country, there has been institutional progress regarding the IAS issue and the development of a national policy, which has made it possible to guide management work across the country (Brasil 2019). More specifically, the Fernando de Noronha Archipelago is part of the MPAs managed by the Núcleo de Gestão Integrada de Fernando de Noronha (ICMBio-Noronha), and by state and regional institutions that deal with territorial use of the urban area. With the arrival of the lionfish in Brazil (Ferreira et al. 2015), it was possible to implement various long-term actions for the management of this IAS with a focus on Fernando de Noronha, including prevention, containment, and mitigation to control the population. Since the first record (Luiz Jr. et al. 2021), the lionfish invasion has been monitored locally with the cooperation of several stakeholders, such as Dive Centers, non-governmental institutes, universities, and local community. Thus, this novel research contributes as a considerable example of IAS management in the South Atlantic. Here we discuss the context of the lionfish invasion in the Fernando de Noronha Archipelago, including how we applied the management strategy to mitigate it. We aimed to 1) report on the history of the strategy applied to lionfish management and 2) describe the temporal and spatial variation recorded by multi-staff agents as a result of the presence of this new population in an isolated South Atlantic archipelago. In this context, this is the first article to discuss long-term (2020–2025) information on lionfish ( Pterois volitans ) management in the South Atlantic, providing new data on ecological drivers (e.g. currents, depth, substrate) on occurrence, lionfish population density and size. Materials and methods Study area Fernando de Noronha (c.03°52′S; 32°25′W) is a Brazilian archipelago located 345 km northeast of the state of Rio Grande do Norte, on the mainland, and about 145 km from the nearest territory, Atol das Rocas (c.03°15’S, 33°40’W). It is the largest oceanic island in Brazil with 21 islands (26 km² of total area; 17.6 km long and 60 km perimeter), raised to a depth of 4,000 m and an altitude of 323 m from its volcanic origin in the Fernando de Noronha Submerged Chain, a group of mountains aligned with the continental shelf (Rocha 1995; Linsker 2003; Teixeira et al. 2003). Classified as an Awi Koppen tropical climate, the region is also under the influence of easterly surface winds that affect conditions mainly on windward cost to the south-east, in addition to swell conditions from leeward coast to the north-west between December to March (Eston et al. 1986; Gouveia et al. 2009; Mello et al. 2023). As for the marine environment, the reef consists of a consolidated substrate (i.e. rocky reef) up to 30 m, covered predominantly by turf, coralline, and macroalgae (Eston et al. 1986; Krajewski and Floeter 2011; Matheus et al. 2019). The deeper zones have a mostly sandy composition with the presence of sponges and rhodolith beds, as the coral cover increases to ~ 5–20% along a depth gradient (Amado-Filho et al. 2012; Matheus et al. 2019). Fernando de Noronha has around 250 recorded fish species, including endemics and threatened ones (Floeter et al. 2008; Pinheiro et al. 2018; Pimentel et al. 2020), such Rocas gregory Stegastes rocasensis , Noronha wrasse Thalassoma noronhanum and the parrotfish Sparisoma sp., genres already verified in the diet composition items of the lionfish in non-native areas (Eddy et al. 2016; Savva et al. 2020; Murillo-Pérez et al. 2021; del Río et al. 2022) and also in stomach context analysis from lionfish from Fernando de Noronha Archipelago, Brazil (Xavier et al. pers. com.). It is the only oceanic island in Brazil with permanent human community, with increasing c. 4,000 people (IBGE 2023), and the Protected Areas in the archipelago's territory include two distinct uses: the Environmental Protected Area (APA; sustainable use; MMA, 2017), and the National Marine Park of Fernando de Noronha (PARNA; strict protection; IBAMA 1990). This last demarks the delimitation over the 50-meter isobath in the marine area. Recreational SCUBA divers visit the two protected areas at different sites, most at depths of up to 20 m, and eventually use open circuits, rebreathers or technical diving for access deeper and longer dives. On-board operations are conducted by Diving Centers (DC), three of which are authorized for commercial guided tourism in the PARNA (numbered as DC1–3), in addition to one focused on lionfish management in this same area and APA (DC4). Five Image Companies (IC) are authorized to accompany DC operations, focusing on selling photo and video media. Diving operations are organized usually with two dives per shift and company (morning and afternoon, with night dives less frequently) with c.50 minutes per site. This results in an average of ten boat operations and 20 active groups of divers per day in the Archipelago, reaching c.10,000 dives per year. Coastal SCUBA diving is also allowed daily in the port region, and freediving is one of the commonest tourism activities in shallow areas close to the main island. Fernando de Noronha has a history involving terrestrial IAS (see Micheletti et al. 2020), but there have never been reports of IAS in the marine environment. In terms of invasive expansion, the lionfish was introduced in Florida in the 1980s and has been increasing its occupation of the South Atlantic (Schofield 2010; Johnston and Purkis 2011). In December of 2020, the species arrived in the Fernando de Noronha Archipelago (Luiz Jr. et al. 2021) and is moving on to other coastal areas (Soares et al. 2022; Maggioni et al. 2023). Strategic alignment according to legal terms In dealing with a new IAS invasion, we report on the application of the management strategy, including all steps for the arrival of the lionfish in Fernando de Noronha archipelago. Preventive works were carried out before the invasion by implementing educational initiatives in the Fernando de Noronha archipelago. After the first sighting of lionfish on the island, we follow the proceedings based on Ordinance No. 19/2025 (Brazil, 2025), a legal instrument that regulates nationally the prevention and management actions for the control and eradication of IAS in Natural Protected Areas and their surroundings. The standardized technical document is officialized as the “Management Project” (MP), which was carried out by the ICMBio-Noronha to its validation by higher authorities. This includes standard actions, dissemination and education activities, specimen destination, adequate training and license for the capture of individuals, and the permission of local agents not directly linked to the local environmental agency was able to participate spontaneously, via a consent term (“Adhesion Term”), to promote the lionfish management. By the collaborative effort of multiple institutions, we call as “multi-staff” all the stakeholders and team members who contributed to the actions of the management process. Control based on multi-staff integration By consolidating the management strategies, local agents, including DCs, ICs and autonomous (e.g. visitors guide, fishers and residents) staff, were training in the characteristics and methods for individual captures and inscribed to follow specific rules regarding lionfish management. Especially when dive operations was targeting a higher effort to lionfish captures, the following topics were clarified: 1. Diving with a focus on lionfish detection (e.g. lionfish-focused search; Green, 2012); 2. Keeping the dive profile suitable for the methodology, ensuring a proper scanning of the substrate; 3. Carefully observing caves, holes and places with less light conducting right to not impact substrate (i.e. Krieger et al. 2013; Giglio et al. 2020); 4. Paying attention to safety regulations and the risk of accidents involving lionfish; and 5. In the event of a sighting, the person in charge would give an audible or visual signal so that the group could record images and follow with the lionfish capture. Lionfish has a low displacement rate, but they can change location in a short period of time (Jud et al. 2015; Bacheler et al. 2015; Tamburello and Côté 2015). Thus, we instructed all local agents to take photos of individuals and reference them as best they could. Only properly trained and authorized divers captured the lionfish individuals with appropriate equipment: a 90 cm Hawaiian Sling harpoon with three to five barbs and the containment PVC tube (standard size of c.15 cm diameter x c.60 cm length) for safely storing captured lionfish. The kit stayed on the boat during all diving operations or taken together for freediving occasions, besides thermoses with hot water or a heat pack on field trips, in case of contact with the lionfish’s poisonous spines. Therefore, divers reported biotic and abiotic parameters based on the dive computer and personal perception, sending each record through an online form (Table 1 ). All information and specimens were forwarded to the ICMBio-Noronha team for standardization in data input. Thus, we collected biometrics of weight (g) and total length (cm), identifying individuals with numbered plastic tags or graphite on paper records to froze and later send them to other institutions that undertake specific research (e.g. diet and genetics). Table 1 Standard information for local agents to fill in when recording invasive lionfish, sectioned into sightings and captures, followed by a characterization of the site and biotic and abiotic factors related to recorded individuals in the Fernando de Noronha Archipelago (South Atlantic, Brazil). Nº Information 1 Sighting date: 2 Sighting time: 3 Who sighted (first and last name): 4 How many individuals were sighted: 5 Capture date: 6 Capture time: 7 Who captured it (first and last name): 8 How many individuals were captured: 9 Equipment used (harpoon or another accessory): 10 Institute/Diving Center: 11 Dive site/reference: 12 Depth: 13 Water temperature: 14 Current (weak, moderate, or strong): 15 Substrate (sand, algae, corals, or artificial): 17 Fish behavior (rest, or swin): 18 Estimated size ( 20 cm): 19 Other details: Post-fieldwork processing for data analysis Lionfish records were treated as a) captures, being the removal of the individual from the environment using some support instrument; b) sightings, being field observations without capture; and c) suspicions, being inaccurate sightings of lionfish by amateur audiences with no proven identification experience. We do not rule out registering suspicions because this was a preventive and management emergency action. We standardized two operation types of targets to the control of the local invasion: 1) opportunistic records during daily activities, as commercial tourist SCUBA diving, fisheries and recreational visitation (hereafter “Routine”) and 2) high effort (i.e. Underwater Visual Transects) for lionfish records during SCUBA or freediving (hereafter “Monitoring”). Group size and time during dives varies widely and was not a predictor between ratings. Therefore, we standardized all localities with lionfish records with a total area of 1.5 hectares (ha; c. 15,000 m²; adapted from Luiz Jr., 2009) to evaluate lionfish density per point based on recorded events over time. Points visited in the same month were corrected as a single recorded event, to avoid underestimating lionfish density. We trademarked 20 m for categorizing all recorded events as shallow ( 20 m) waters, by considering the profile usually performed in dive points by local agents and limits of diving certifications. Population estimate values were evaluated including the total number of records during the study period, by applying the intrinsic growth rate ( r ): \(\:r=\text{l}\text{n}\left[\left({N}_{t2}/{N}_{t1}\right)\right]/(t2-t1)\) , and increment rate ( λ ): \(\:\lambda\:={e}^{r}\) (e.g., Margalef 1998), where \(\:\text{l}\text{n}\) is the logarithm of the fraction between the final ( \(\:{N}_{t2}\) ) and initial ( \(\:{N}_{t1}\) ) population size, and \(\:e\) is the base of the natural logarithm (~ 2.718). We analyzed the length-weight (L-W) relationship using the allometric growth function \(\:W=a\times\:{\:L}^{\:b}\) and later the body condition factor (Fulton’s K) through \(\:K=(W/{L}^{3})\times\:100\) . To better characterize the lionfish detection scenario in the archipelago region, we compared the frequency of records and their respective information. We use multiple regression to check the relation between density by time and depth, factorial ANOVA to relate records between the operation types (routine/monitoring) and depth, as the same for density per month to check inter-annual variation. To compare the complementary data provided by the records, we use Chi-Square with expected proportion tests with no difference between categories, including divers' perceptions about the substrate (benthic organism or soil backgrounded with the fish), marine current (drift speed at each dive point) and fish behavior (individual activity status). Generalized Linear Model (GLM) was applied to compare protected areas, sea regions and body conditions on depth and time. Detected sites with more than 3 lionfish recorded events were evaluated to compare the effect of management efforts on density by time, using the slope coefficient of linear regression to classify decrease ( 0) trends. To spatially analyze of lionfish distribution in the Fernando de Noronha Archipelago we estimate the most explored areas for each bathymetry features (20, 50, 100 e 500 m depth) categorizing five classes (0–70 m as well explored, 71–100 few explored, 101–150 inferred, 151–300 distant, and 301–4000 very distant) by referring a Euclidian Distance raster. In order to predict the size of the fish population in this area, we considered the maximum density values for each location with a recorded event and used the geostatistical interpolation method by ordinary kriging. The values were transformed using the natural logarithm (ln) for better adjustment of the semivariogram Gaussian model. Subsequently, the data was reprocessed for spatial analysis of the species density distribution. The population estimate was based on the sum of the estimated density values in all cells (100 m²) within the study area. We used QGis v3.40 ('r.grow.distance' and 'Smart-Map' tools) for the spatial analyses and map output and RStudio v.2023.04.1 for the statistical tests and visual graphics (α = 0.05). Results Reporting on management strategy processes We characterized five main steps based on Brazilian policy scenario for IAS management protocol (Ordinance No. 19/2025), including the period before the lionfish invasion begins to the stabilization of its management strategy: 1) prevention about IAS; 2) IAS record; 3) preparation and submission of MP; 4) evaluation and MP approval; and 5) upon approval, implementation of the “Adhesion Term” in order to regulate the management actions by local agents (Table 2 ). All the actions for the lionfish in Fernando de Noronha were taken by the engagement of multiple institutions to procedure with its management (Fig. 1 ). Before the arrival of the lionfish in the Fernando de Noronha Archipelago, previous publications focused on prevention for Brazil (e.g. ICMBio 2019) and relevant actions were also made locally in 2018–2019 by Projeto Conservação Recifal (PCR; https://conservacaorecifal.com/ ), focusing on educational actions and training the government agencies, divers, fishermen, residents, and tourists, about the lionfish threat as IAS (Fig. 1 A). The first lionfish was detected on 20 December 2020, supported by the Universidade Federal Fluminense (UFF) with DC4 (under authorization SISBIO 41327; Luiz Jr. et al. 2021). To enabled rapid communication and first actions with local agents (mainly the DCs), the “Emergency strategy for lionfish management in the Fernando de Noronha Archipelago” (ICMBio 2021) was consolidated by the ICMBio-Noronha on 30 March 2021. This strategic protocol had resulted in the initial training and lionfish captures (n = 25), while the MP was in progress. The preparation and submission of MP was made by ICMBio-Noronha on 6 October 2021, which was ratified 30 days after submission. This made up the ongoing instructional procedure that regulates the multi-staff agents (108 members and all the four DC which works onboard) to act via “Adhesion Term” on lionfish management in Fernando de Noronha. Since this period, it was totalized 2,660 lionfish records in all Archipelago. In addition, on 10 September 2024, Ordinance No. 2,761 revised the Management Plan of the Fernando de Noronha Marine National Park (i.e. IBAMA 1990). This regulatory amendment enabled IAS control activities associated with tourism-related operations throughout legally and autonomously participate of dive operations, providing a complementary mechanism to support the financial sustainability of ongoing control efforts. Table 2 Process for structuring an IAS Management Project (MP), according to the Brazilian policy scenario (Ordinance No. 6/2019), compared with the lionfish invasion in the Fernando de Noronha Archipelago (this paper) and the management effort of fieldwork over time. Step 1 Step 2 Step 3 Step 4 Step 5 Total Management process based on Brazilian guidelines * Prevention about IAS IAS record Preparation and submission of MP Evaluation and MP approval Adhesion Term for local agents Total Lionfish case in Fernando de Noronha 2018–2019 20 December 2020 6 October 2021 5 November 2021 18 November 2021 (ongoing) 20 December 2020 to 30 September 2025 (ongoing) Duration of the process - - 291 days (~ 11 months) 321 days (~ 12 months) 334 days (~ 12 months) 1,746 days (~ 58 months) Number of lionfish records 0 1 20 25 28 2,660 Validated document for management - Emergency strategy (ICMBio 2021) Emergency strategy (ICMBio 2021) Emergency strategy (ICMBio 2021) Management Project (ongoing) - Support lionfish management was performed between the multi-staff institutions covered by specific roles for decision-making from local to international level (Fig. 2 ). Permission and institutional orientation through MP alignment were provided on a national scale by the high instance of national government represented by Coordination of Invasive Alien Species Management (ICMBio-CMEEI). From a regional scale, technical guidance, planning and specific research (including specimen destination) was subsided by Non-governmental organization PCR and Universities, such as Universidade Federal Fluminense (UFF), Universidade Federal do Rio Grande de Norte (UFRN), Universidade Federal de Pernambuco (UFPE), Universidade Federal de Alagoas (UFAL), and Universidade Federal do Ceará (UFC), besides network engagement for planning actions by Pernambuco State Environment Agency (SEMAS/CPRH; Fig. 1 C). International support included key capacity building through inter-institutional exchanges with experts (represented by PB) of Bonaire’s Environmental Agency, the Stichting Nationale Parken Bonaire (STINAPA; Fig. 1 B), as well as specific research of California Academy of Sciences (CAS). Thus, the ICMBio-Noronha assumed a central role to coordinate multiple institutions, overall, to maintain regular training sessions, processing data and specimens and stimulate lionfish management with local DCs and ICs, which provide most lionfish records during routine and monitoring operations (n = 2,614; 98.3%; Fig. D–F). Results outcome by multi-staff integration During the period from 20 December 2020 to 30 September 2025, there were registered 2,404 captures, 251 sightings and 5 suspicions (N = 2,660) of lionfish in the Fernando de Noronha Archipelago, featuring an exponential increase of 345.9% in its population over this period ( r = 1.495; λ = 4.459; Fig. 3 A). It was observed that the highest value reached 299 records in September 2025 (292 captures), followed by June 2025 (n = 213; 208captures) and April 2025 (n = 126; 125captures). After the first record in December 2020, there were no detections for six months of lionfish individuals, when the second individual was captured in June 2021, presenting a continuous increase every month since then (45.1 ± 56.3 records/month). Through 52 months, records amounted to 407 events (Fig. 3 B), gradually increasing each year until they peak in 2025 between shallow (n = 184 records; 44 events) and deep waters (n = 1030 records; 85 events). Regarding inter-annual variations, lionfish densities have not varied significantly between months (F 11,73 = 0.526; p = 0.874). Moreover, we could check that densities were related positively with time and deeper records (R 2 = 0.521; p < 0.001), by presenting higher densities in deep waters (8.2 ± 3.9 individuals/ha) than compared with shallow regions (2.5 ± 2.1 individuals/ha) in recent years (t 2025 ). Thus, the general scenario of lionfish invasion is estimated at 3.9 individuals/ha for the Fernando de Noronha Archipelago. Depth gradient showed a normal distribution (33.0 ± 12.1 m), counting with more individuals between 20 to 40 m (n = 1,593; 62.3%) and less records in shallower (34 records above 10 m; min.: 0.5 m) and deeper (130 records under 50 m; max.: 95 m) areas. Records were related significantly with deep waters (F 2,1726 = 16.010; p < 0.001) and monitoring operations (F 1,4937 = 45.795; p < 0.001), which presented major values and variation for the number of records (6.9 ± 9.3 ind./ha) than opportunistic routine records (1.8 ± 2.1 ind./ha). In addition, SCUBA diving was the most collaborative activity for field detection in all Archipelago (n = 2,634; 99%), with few records made up during freediving (n = 24; 0.9%). A single catch by a fisherman was verified and one case when the lionfish was filmed almost at the surface by a person above water at the Port (Table 3 ). Thus, recorded lionfish were mainly performed by Dive Instructors (n = 2,597; 97.6%), and, less representative, by other agents, such as camera divers, tourists, fishers, and residents (n = 63; 2.3%). Great effort was realized by DC4 (n = 2,278; 85.6%), mainly related to focused operations on monitoring deep areas in the Archipelago, followed by those carrying out largely routine operations by DC1–3 and ICs (n = 336; 12.6%). Although not always provided, complemented information was noted several occasions by multi-staff registers, regarding most records for I) recording time (n [morning] = 584; 53.4%), II) marine current (n [weak] = 1911; 91.3%), III) substrate composition (n [coral] = 1,012; 49.6%), and IV) fish behavior (n [rest] = 1963; 99.4%). Table 3 Data parameters on lionfish records in the Fernando de Noronha Archipelago gathered by local agents, including the operation type, related activity, local agents, Dive Centers (DC), recording time, marine current, substrate composition, fish behavior, and Protected Natural Areas, by categories of suspect, sighting, and captures. Each section is accompanied by the Chi-Square result and its significance level (* represents the significant category). Categories Suspect Sighting Captures Total Method (χ 2 = 1295.0; p < 0.001; N = 2,660) Monitoring * 0 105 2,153 2,258 (84.9%) Routine 5 146 251 402 (15.1%) Activity (χ 2 = 7773.9; p < 0.001; N = 2,660) SCUBA diving * 0 242 2,392 2,634 (99%) Freediving 5 8 11 24 (0.9%) Fishing 0 0 1 1 (0,04%) Above water 0 1 0 1 (0.04%) Local agents (χ 2 = 10021.0; p < 0.001; N = 2,660) Dive Instructors * 0 216 2,381 2,597 (97.6%) Camera divers 0 16 3 19 (0.7%) Tourists 3 3 0 6 (0.2%) Fishers 0 0 1 1 (0.04%) Others 2 16 19 37 (1.4%) Dive Centers (χ 2 = 9142.6; p < 0.001; N = 2,660) DC1 0 48 27 75 (2.8%) DC2 0 41 130 171 (6.4%) DC3 0 17 61 78 (2.9%) DC4 * 0 117 2,161 2,278 (85.6%) ICs 0 10 2 12 (0.5%) Others 5 18 23 46 (1.7%) Recording time (χ 2 = 6.8; p = 0.009; N = 1,086) Morning * 3 84 499 586 (54.0%) Afternoon 2 36 462 500 (46.0%) Marine current (χ 2 = 3186.3; p < 0.001; N = 2,099) Weak * 5 156 1,754 1,915 (91.2%) Moderate 0 14 158 175 (8.3%) Strong 0 0 9 9 (0.4%) Substrate composition (χ 2 = 1964.7; p < 0.001; N = 2,039) Sand 0 17 63 80 (3.0%) Algae 0 28 446 474 (17.8%) Coral * 0 85 927 1,012 (38.1%) Rock 2 46 242 290 (10.9%) Artificial 0 1 43 44 (1.7%) Porifera 0 6 133 139 (5.2%) Fish behavior (χ 2 = 1927.3; p < 0.001; N = 1,975) Rest * 1 162 1,797 1,963 (73.8%) Swin 0 3 9 12 (0.5%) Protected Areas (χ 2 = 320.9; p < 0.001; N = 2,660) APA 1 73 794 868 (32.6%) PARNA * 4 178 1,610 1,792 (67.4%) Total 5 (0.2%) 251 (9.4%) 2,404 (90.4%) 2,660 (100%) Individuals generally average 23.17 ± 6.7 cm of total length, recording smaller (8.1 cm) to bigger (46.0 cm) detections. Weight remained between 238.2 ± 241.6 g, ranging from 7.0 to 1670.0 g. There was a significant difference over the years for both parameters, with higher values for size (F 5, 1275 = 18.594; p < 0.001) and weight (F 5, 1275 = 10.986; p < 0.001) in recent years. The L-W relationship showed a strong association (R 2 = 0.96; p < 0.001) and high coefficient (b = 3.22), pointing to positive allometric growth and larger individuals presenting proportionally more body mass in relation to length (Fig. 4 A). The body condition factor reveals high physiological condition (Fulton’s K = 1.43). The effect of time on the condition factor K was significant (F 5, 1275 = 16.76, p < 0.001), indicating that the body condition of the population varied between years (Fig. 4 B). Despite the wide variation, body size was significantly related to depth, with higher averages (K: 1.38 [deep] > 1.31 [shallow] ) for individuals at greater depths (F 1, 606 = 7.028; p = 0.008). Mult-staff effort totalized 116 localities, which represented c.610.50 ha throughout the 407 recorded events during this study period. Lionfish individuals were recorded in both protected areas of PARNA (n = 1,792; 67.4%) and APA (n = 868; 32.6%), besides regions of Leeward (n = 2,289; 86.2%) and Windward (n = 367; 13.8%) Sea regions. Species density was significantly greater in the PARNA (5.2 ± 8.7 ind./ha) than in the APA (3.7 ± 4.9 ind./ha) areas (β = 0.338; p < 0.001), and for leeward (5.0 ± 8.2 ind./ha) than in windward (2.9 ± 3.1 ind./ha) sea regions (β = − 0.522; p < 0.001). Most of recorded events showed lower densities (n = 305; 78.6%), ranging from 1 to 5 individuals per point. Higher densities were found in three unusual points located in Leeward Sea and in the PARNA, counting respectively with 90 (60 ind./ha), 80 (53 ind./ha), and 63 (42 ind./ha) lionfish records in single visits to the dive sites. Official dive sites (i.e. authorized for commercial visitation) were surveyed in 18 localities and represent 349 lionfish records. These points presented lower densities in general (2.1 ± 3.3 ind./ha), although high detections were checked in Cagarras Fundas (n = 101 records), Laje Dois Irmãos (n = 81), and Cabeço da Sapata (n = 59) over the study period. The effect of field management proved to be effective in maintaining a low temporal trend (slope coefficient median = 0.02) throughout the evaluated period (Fig. 5 ). Most of repeatedly visited sites (n [S1–S45] = 45; 6.51 ± 4.70 visits/site) showed stable trends (n = 30; a ≈ 0) with low density averages (2.01 ± 1.68 ind./ha), although there has been an overall average increase (0.10 ± 0.46 coefficient units) and some locations (n = 10; 22%) recorded pronounced growth ( a > 0.13). Euclidean distance map presented a general value of 1,364 ± 956 m (Fig. 6 ), pointing to 17,331.98 ha (89.4%) of entire study area (0–500 m depth) as very distance regions (Table 4 ). Well explored areas (181.51 ha; 0.9%) was checked over 20 m (98.91 ha; 54.1%), followed by 21 m to 50 m (68.82 ha; 37.9.0%), 51 m to 100 m (10.87 ha; 6.0%), and 500 m to 100 m (2.91 ha; 1.6%), besides few explored areas (172.13 ha; 0.9%), which totalizing both 353.64 ha (1.8%) of most recognized areas surround the Archipelago. The adjusted variogram model showed a good predictive capacity (RMSE = 0.414; R² = 0.661) and positive spatial autocorrelation by presenting local density clusters through interpolation analysis (Moran's I = 0.161; p < 0.001). The kriging map showed the presence of areas of higher density (above 20 ind./ha) concentrated in specific sectors of leeward sea, with a decreasing gradient toward the remote areas (Fig. 7 ). Thus, the population estimate using this analysis reached 75,742 (CI 95% 56,202–95,372) individuals and lionfish density of 3.90 (CI 95% 2.89–4.91) ind./ha in Fernando de Noronha Archipelago. Table 4 Estimated lionfish sampling areas (ha; percentage) into five effort classes (well explored: 0–70 m; few explored: 71–100 m; inferred: 101–150 m; distant: 151–300 m; very distant: 301–4000 m) and between the sectors of the isobath lines from 0 to 20, 21 to 250, 51 to 100, and 101 to 500 m depth. Effort classes Isobaths classes (m) Total 0–20 21–50 51–100 101–500 Well explored 98.91 (54.5%) 68.82 (37.9%) 10.87 (6.0%) 2.91 (1.6%) 181.51 (0.9%) Few explored 86.23 (50.1%) 71.97 (41.8%) 11.23 (6.5%) 2.7 (1.6%) 172.13 (0.9%) Inferred 168.86 (45.6%) 167.47 (45.2%) 26.86 (7.3%) 6.96 (1.9%) 370.15 (1.9%) Distant 476.34 (36.1%) 661.01 (50.0%) 145.86 (11.0%) 37.52 (2.8%) 1,320.73 (6.8%) Very distant 1,151.29 (6.6%) 8,047.75 (46.4%) 3,751.19 (21.6%) 4,381.75 (25.3%) 17,331.98 (89.4%) Total 1,981.63 (10.2%) 9,017.02 (46.5%) 3,946.01 (20.4%) 4,431.84 (22.9%) 19,376.5 (100%) Discussion The management strategy presented herein demonstrates that the use of legal and protocolary adoptions have efficiently stabilized and outlined the lionfish control through standardized actions in the Fernando de Noronha Archipelago. Combining multi-staff actions based on international and national protocols with a strong local field effort, it was possible to designate specific roles covering better decision-making. Consequently, these actions generated positive results for understanding lionfish population and promoting a protocol for local agents, which was reflected in the number of records and practical management in the South Atlantic. Even considering the high population level locally, we could check that the effort in several sites was effective to control lionfish density throughout the time, including a decrease over time in some areas. Those results reflect the main strategy to be maintained in areas affected by lionfish invasion, even as a form of local control, in addition to encouraging management of the individuals captures (Green et al. 2014; Bogdanof et al. 2021). In comparison to other locations, we observed that similar protocols have also been effective for more complex and intense lionfish invasions, such as Caribbean islands and Mediterranean Sea (Dahl et al. 2019; Ulman et al. 2022). We highlight that the management strategy for lionfish control in FNA constitutes a model to be applied in different locations, which must be adapted according to the scale to be followed and its regulations. Lionfish management in Fernando de Noronha showed that SCUBA diving and the active participation of dive instructors were key factors in controlling the species. These results are like other regions that have shown the integration of various teams in lionfish removal actions (Anderson et al. 2017). This involved the entire time series related to the detection of this IAS, as observed since the first record of the species in 2020. All steps of the management project were designed to include divers in effective control, given that this activity is one of the most practiced in the archipelago and that the areas most accessed by citizen scientist divers are shallow coastal subtidal areas, with depths < 40 m (Thiel et al. 2014). Freediving participation was also significant, although the number of records was lower. This may be related to the higher density in deep areas compared to shallow waters. Furthermore, there was a significant increase in the number of individuals below 20 m depth, especially in 2025, noting that the density trend in shallow waters can increase more slowly over time. This pattern is recognized for lionfish invasion processes, and their colonization may differ between depth levels, with levels in shallow and deep waters becoming similar over time (Airey et al. 2023). Although we have observed stable trends in several areas, density trends at some sites and the cumulative curve show that lionfish occupation in general is progressing rapidly and is probably greater than management efforts can support. The pattern in IAS begins with low-density introduction and establishment levels and exponentially increasing density and population growth rates (Sakai et al. 2001; Dahl et al. 2019), a similar pattern observed in these early years of lionfish recognition in our study site. In addition, our work focuses mainly on relative effort measures, based on the number of detections provided in each record reported by local agents. It should be noted that some sites receive continuous visits, such as official commercial dive sites and shallower areas frequented by freedivers, which is considered a significant effort. However, by checking nested clusters for the spatial distribution of lionfish population in Fernando de Noronha, we can infer that other biological variables may have a stronger influence on distribution throughout the archipelago. Lionfish populations are denser in deeper waters, related to the range of habitat diversity, main areas of population recruitment due to site fidelity, influence of light penetration, as well as lower pressure from the removal of individuals, with the dispersion of individuals aggregating from deep to shallow waters (Biggs and Olden 2011; Nuttal 2014; Airey et al. 2023). Despite the effort undertaken in this study, there was less frequent diving in mesophotic regions, which presented greater technical difficulty and consequently a lag in the effort to estimate individuals. Previous studies have demonstrated that lionfish populations frequently reach higher densities and comprise larger individuals within marine protected areas (MPAs), when compared to adjacent non-protected zones (Morris et al. 2011; de León et al. 2013; Andradi-Brown et al. 2017; Kleitou et al. 2024). In this regard, lionfish currently occupy 33% of Brazil’s MPAs and are likely to spread to an additional 25 protected areas (c.60% of MPAs) within the next decade, while regular removal programs are absent in most of them (Soares et al. 2025). Such findings emphasize the challenge of managing lionfish in protected areas, where no-take zones should be specially treated with IAS removal through management strategies because restrictions on extractive activities create favorable conditions for invasive populations to persist. As observed, lionfish distribution in Fernando de Noronha is not specifically related to the boundaries of protected areas. High and low-density points were found in both visited and unvisited areas, which reinforces that the sampling effort applied for management should be applied more systematically. In this case, the interpretations obtained to affect population control can be better grounded in ecological factors and through planned mapping. This also reinforces decision-making regarding invasion areas, showing that continuous monitoring and local agents training, especially diving instructors, should be done more frequently. Considering the negative impacts caused by the intensity of recreational diving in various locations (Giglio et al. 2020; 2025), caution and good planning are needed to open new sites, especially in protected areas (Soares et al. 2023). We therefore encourage lionfish management actions to be carried out with the support of planned strategies. For instance, considering sea conditions and diving safety, it would be optimal to apply a grid of plots over the marine region of the archipelago and sampling these locations through monitoring operations, by applying focused census and proper removal of lionfish. The role of diving operators proved to be highly relevant in ensuring that IAS control was more successful. It was observed that one Dive Center (DC4) made a great effort regarding the number of records, including the lionfish sightings and, mainly, captures. This was related that DC4 maintains continuous operations with a greater focus on lionfish control, being recognized and legally authorized to operate in the protected areas around the archipelago. This strategy was well interpreted in terms of management strategy due to the difference between diving operation methods. Operations in the PARNA is performed historically before lionfish invasion by three specific dive centers (DC1–3), as this is a region that requires greater control due to its environmental sensitivity. Considering that the recreative commercial model of these operations was focused on customer service and recreational diving safety, we call them routine operations, which also contributed in an important way to the mapping of records in shallow waters and at official sites in PARNA. Thus, it became quite strategic to regulate the format of operations focused on lionfish capture, referred to here as monitoring operations, resulting in greater recognition of sites and increasing the number of records. Similarly, these activities must be well coordinated so as not to cause overlaps between groups and greater impact on diving spots (Luiz Jr. 2009). In addition, the routine operations are mainly carried out in shallow waters, which have been ecologically linked to lower lionfish detection rates. By observing lionfish increase over time at these sites, it is crucial to conduct more operations focused on species control, balancing them with recreational operations. Local communication was also a distinctive factor in making IAS control practical. Population control and action protocols tend to be better managed and controlled in places with a smaller territorial scale, due to the possibility of applying management efforts in the field and communication between local agents in a simpler and more effective way (Earp and Liconti 2020). This reflects in better data updating and, thus, in the understanding of parameters related to the lionfish population (Kelly et al. 2020). We consider that the reporting of records via virtual app was effective in proceeding with the analyses applied here, which should be further encouraged, so that the number of registered individuals could be disclosed faster by interested media and all teams involved. Regarding the management permission process, despite the extensive bureaucracy of Brazilian public processes in general, this procedure was carried out with reasonable promptness. Since the first lionfish was detected on 20 December 2020, an Emergency Strategy was published in the following month to guide the first actions to multi-staff integration and practice lionfish control in the archipelago (ICMBio 2021). This was carried out by ICMBio-Noronha, with the support of ICMBio-CMEEI, to begin jointly within a month after the presentation of the MP. It is important to note that the steps followed for the approval of lionfish management project were reinforced by proper evaluation, following standardized actions for each location and which were in accordance with the national protocols (ICMBio 2019). Such procedures are useful for developing better strategies and facilitating multi-staff adhesion, resulting in the obtaining of explanatory ecological variables for IAS control. For the specimens found in Fernando de Noronha, a high physiological condition was observed and the L-W relationship, indicating high consistency between measurements. The allometric coefficient was positive, characterizing that the growth of larger individuals presents proportionally more body mass in relation to length. This scenario indicates that the lionfish population in this region exhibits growth typical of species with good energy supply and favorable body condition (Edwards et al. 2014; Chávez-López et al. 2025). These factors are associated with good nutritional condition, an environment with good resource availability, and the absence of persistent energy stress (del Río et al. 2023). Substrate composition in Fernando de Noronha presented a variety of environments, in reference to the field records, with greater detection for epibenthic organisms related to corals, but also for algae and porifera, in addition to artificial structures, such as shipwrecks and anchors. Lionfish individuals are associated with a diverse set of habitats including different conditions of visibility, lightness, depth and environments (Schofield 2010). Fish behavior was noted with greater frequency for fish in resting behavior, which is consistent with the well-known low mobile behavior of lionfish (Jud and Layman 2012). In addition, we also encourage efforts to detect seasonal patterns over time. Despite the temporal series, the initial invasion of lionfish in archipelago did not allow for a relevant sampling effort to detect differences between seasons or relationships with sea conditions, which may be better indicated with a greater establishment of the species. Our work corroborates with Soares et al. (2023), especially regarding collaborative network of actions for raising public awareness and developing education programs. We have provided an important baseline on the invasion of this species in new locations in South Atlantic, in addition to adequately addressing the procedures for adapting and integrating multiple teams. The local reality of Fernando de Noronha is composed of singular features, considering it is an isolated and small territory. The invasion of lionfish continues to advance to other locations in Brazil, and other difficulties may influence the control of this species, such as the frequency of diving, standardized training of multi-staff, and specific regulations. We add that each location could be delimited, based on protected areas boundary or municipalities, and communities. This makes it possible to understand local realities and apply standardized protocols through regulations. Also, new methods of control must be employed, especially in waters that are not suitable for diving, where it is possible to apply trapping techniques, such as pots and other fishing techniques (Pitt and Trott 2015). That scenario is still lacking and includes improving control methods in mesophotic and rough marine environments (Gress et al. 2017), such as the leeward sea in Fernando de Noronha. In conclusion, this research can add important perspectives for the development of national or regional regulations (e.g., ICMBio 2019; 2021; CPRH 2024), as well as the integration of international protocols, since the invasion of lionfish depends on an integrated effort (Green 2012). In addition, important advances regarding the use of captured individuals, such as gastronomic consumption, can be made regularly as a form of control. These actions should always be reinforced by educational activities, as we presented that open communication and multi-staff integration have become crucial tools for promoting an adequate strategy for lionfish control. Declarations Conflict of interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Author contributions The project was conceptualized by RA, CBPES, LPSS, PPC, TEC and TCSG. All authors contributed to the project administration, resources, and data collection. Data analyses were performed by Lucas Penna Soares Santos. The first draft of the manuscript was written by Clara Buck Pereira do Eirado Silva and Lucas Penna Soares Santos. All authors commented on previous versions of the manuscript. All authors read and approved of the final manuscript. Acknowledgements This work forms the coordination of research and management of ICMBio-Noronha, represented by R. Araújo. Ethical permissions were supported by ICMBio-CMEEI (Management Project/Permission n. 10/2021 SEI n. 9919898). We thank all local agents who worked collaboratively and contributed to the lionfish management strategy in Fernando de Noronha, especially the dive instructors M. Wilson (Mike), R. Rocha (Beto), A. Gonzaga, A. Pereira (Babu), F. da Costa (Lula), J. Huss, and L. Lopes. A special strengthening was provided by Sea Paradise, represented by F. Rodrigues, the Dive Centers of Águas Claras, Atlantis Divers, Noronha Diver, Mar de Noronha, Maravista Mergulho, as well as the Image Companies of All Angle, Barracudas, Ciliares, Hidrosfera, and Natureza Viva. We appreciate all the support provided by STINAPA, especially P. Bertuol, Projeto Conservação Recifal (PCR), particularly P. Cipresso and L. Guilherme, PELD-ILOC, and researchers. SEMAS and CPRH established the network to control lionfish in the state of Pernambuco, with the cooperation of P. Tavares, S. Vieira, and D. Alves. To the ICMBio-Noronha Team, the Administration of the Port, Corpo de Bombeiros, visitors’ guides, boat operators and community of Fernando de Noronha. We thank the reviewers who contributed to this article. Data availability Data will be made available on request. References Airey ME, Fogg AQ, Drew JA (2023) Invasive lionfish dispersal between shallow-and deep-water habitats within coastal Floridian waters. 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Mar Pollut Bull 49:844 − 849. https://doi.org/10.1016/j.marpolbul.2004.05.001 Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 03 Mar, 2026 Reviewers invited by journal 02 Feb, 2026 Editor invited by journal 23 Jan, 2026 Editor assigned by journal 20 Jan, 2026 First submitted to journal 19 Jan, 2026 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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C. Guaitanele","email":"","orcid":"","institution":"Chico Mendes Institute for Biodiversity Conservation: Instituto Chico Mendes de Conservacao da Biodiversidade","correspondingAuthor":false,"prefix":"","firstName":"Carla","middleName":"C. C.","lastName":"Guaitanele","suffix":""},{"id":584308599,"identity":"7d7fc0e7-7f02-4e8a-b882-fd70b332d82a","order_by":18,"name":"Lilian L. Mitiko Hangae","email":"","orcid":"","institution":"Chico Mendes Institute for Biodiversity Conservation: Instituto Chico Mendes de Conservacao da Biodiversidade","correspondingAuthor":false,"prefix":"","firstName":"Lilian","middleName":"L. Mitiko","lastName":"Hangae","suffix":""},{"id":584308600,"identity":"5dcc0e2a-56ec-4374-96a3-dd66f34c4944","order_by":19,"name":"Mario Douglas Fortini","email":"","orcid":"","institution":"Chico Mendes Institute for Biodiversity Conservation: Instituto Chico Mendes de Conservacao da Biodiversidade","correspondingAuthor":false,"prefix":"","firstName":"Mario","middleName":"Douglas","lastName":"Fortini","suffix":""},{"id":584308601,"identity":"a5368dbe-e0c5-4189-ab7c-70023830b6dd","order_by":20,"name":"Ricardo Araújo","email":"","orcid":"","institution":"Chico Mendes Institute for Biodiversity Conservation: Instituto Chico Mendes de Conservacao da Biodiversidade","correspondingAuthor":false,"prefix":"","firstName":"Ricardo","middleName":"","lastName":"Araújo","suffix":""}],"badges":[],"createdAt":"2026-01-17 15:01:02","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8626827/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8626827/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":101898518,"identity":"8b7d1a5d-97ef-4d64-85b2-4a4ed5e5b4ca","added_by":"auto","created_at":"2026-02-04 18:15:38","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":2114342,"visible":true,"origin":"","legend":"\u003cp\u003eLionfish management strategy in Fernando de Noronha Archipelago (Brazil) reported by multiple institutions and actions, as follow: A) educational work with young public, during \u003cem\u003eProjeto Conservação Recifal\u003c/em\u003e campaign to prevention lionfish threats; B) main training for general public supported by the international team of the Stichting Nationale Parken Bonaire (STINAPA); C) action plan for controlling and monitoring lionfish bioinvasion in Pernambuco seminar; D) multi-staff fieldwork with DCs and ICMBio-Noronha integration for lionfish monitoring; E) lionfish capture as result of multi-staff monitoring during SCUBA diving fieldwork and; F) several captures of individuals performed through lionfish-focused operations.\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-8626827/v1/a6a4d501b4b6f79a77c85eda.png"},{"id":101898514,"identity":"3ce6c16c-8642-469f-b51d-192f0649ed35","added_by":"auto","created_at":"2026-02-04 18:15:38","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":253261,"visible":true,"origin":"","legend":"\u003cp\u003eProcess map of the multi-staff contribution to the lionfish management strategy in the Fernando de Noronha Archipelago. Semicircular blocks represent the functions of each institution and their scale scenario, including: A) national (green) support of legal processes for management license by Coordination of Invasive Alien Species Management of Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio-CMEEI); B) local (orange) engagement and major fieldwork provided by Image Companies (IC) and Dive Center (DC 1–4); regional (blue) involvement for technical support, prevention actions, and specimen destination of research institutions of Projeto Conservação Recifal (PCR), Universidade Federal de Pernambuco (UFPE), Universidade Federal do Rio Grande do Norte (UFRN), Universidade Federal do Alagoas (UFAL), Universidade Federal Fluminense (UFF) and Environmental Department of Pernambuco State Government (SEMAS); and international (blue) expertise training support and specific researches with the Stichting Nationale Parken Bonaire (STINAPA) and California Academy of Sciences (CAS). Pie chart shows the records of lionfish (light orange = sightings; dark orange = captures) made by DCs during fieldwork.\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-8626827/v1/5457305eaaa71c496325e027.png"},{"id":101898515,"identity":"d80bd201-024d-4eb7-b096-e4c30ee62b76","added_by":"auto","created_at":"2026-02-04 18:15:38","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":89306,"visible":true,"origin":"","legend":"\u003cp\u003eA) Timeline presenting light orange bars with monthly records and dark orange lined dots with cumulative curve of lionfish in the Fernando de Noronha Archipelago, from December 2020 to September 2025. B) Number of events that recorded lionfish individuals in shallow waters (above 20 m; light orange bars) and deep waters (under 20 m; dark orange bars), as well as average density (records/ha) of lionfish over years, classified between shallow (clear orange dots and light blue line) and deep (dark orange dots and dark blue line) waters.\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-8626827/v1/9d0c6d6e07dacefe77cfa91a.png"},{"id":101943143,"identity":"02f51b13-bfa5-4e07-85dd-0d356a59cd1d","added_by":"auto","created_at":"2026-02-05 09:40:44","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":72148,"visible":true,"origin":"","legend":"\u003cp\u003eBody parameters of lionfish individuals from Fernando de Noronha Archipelago presenting A) growth model from length-weight (L-W) relationship, with upper (total length) and left (weight) internal histogram bars and B) body condition factor (Fulton’s K) throughout years (2020 – 2025).\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-8626827/v1/f51cbcffdf7fb99fedbee83c.png"},{"id":101898520,"identity":"dcedd2cb-817a-4a27-a36a-39d517e6c2f0","added_by":"auto","created_at":"2026-02-04 18:15:38","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":75726,"visible":true,"origin":"","legend":"\u003cp\u003eManagement effort assessed by sites (1–45) that achieved more than three visits during the study period. For each location, we highlight the visits counts (blue horizontal bars), average lionfish density (orange vertical bars) with standard deviation (upper orange whiskers), and slope coefficient of linear regression (black bordered orange dots), which represents temporal trend of decrease (\u0026lt; 0), stable (≈ 0), and increase (\u0026gt; 0) lionfish density.\u003c/p\u003e","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-8626827/v1/3625301fbf99dc53ee35abcf.png"},{"id":101898517,"identity":"bcb896e4-8890-48c4-9696-ae488bae406e","added_by":"auto","created_at":"2026-02-04 18:15:38","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":864390,"visible":true,"origin":"","legend":"\u003cp\u003eLionfish invasion map within a 500 m bathymetry in Fernando de Noronha Archipelago from December 2020 to September 2025, featuring recorded individuals (black dots) and sampling locations with Euclidian distance (in meters), which estimates the application of the sampling effort for the areas.\u003c/p\u003e","description":"","filename":"floatimage6.png","url":"https://assets-eu.researchsquare.com/files/rs-8626827/v1/5ad608c000808206b7986301.png"},{"id":101898519,"identity":"bed6c7b4-13be-4de2-af88-053b9475d099","added_by":"auto","created_at":"2026-02-04 18:15:38","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":775115,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution model of lionfish invasion within 500 m bathymetry in Fernando de Noronha Archipelago, highlighting density interpolation (100 m² per pixel) based on maximum lionfish density per site (shaded gray circles) detected from December 2020 to September 2025.\u003c/p\u003e","description":"","filename":"floatimage7.png","url":"https://assets-eu.researchsquare.com/files/rs-8626827/v1/a8c33f929e5b57058cfa2dae.png"},{"id":101944683,"identity":"b8b62e89-dae2-4914-8e1f-21766baa5598","added_by":"auto","created_at":"2026-02-05 09:53:41","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":5651201,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8626827/v1/489b2747-b252-467f-9459-d8d61598a8fd.pdf"}],"financialInterests":"","formattedTitle":"Management strategy based on multi-staff integration target control of lionfish invasion on a South Atlantic oceanic island","fulltext":[{"header":"Introduction","content":"\u003cp\u003eA challenging and growing topic for the conservation of natural environments is the introduction of invasive alien species (i.e. intentionally or unintentionally introduced, as the same, non-native species; hereafter IAS). IAS complies the second leading cause of species extinction in natural environments and the reason for large-scale environmental and socioeconomic changes (Molnar et al. 2008; Bellard et al. 2016; Vil\u0026agrave; and Hulme 2018; Giakoumi et al. 2019). Although high-cost, strategies to control IAS presented good results on several cases (Giakoumi et al. 2019), treating as management all the actions needed to prevent, containment, and mitigation (Blackburn et al. 2011). In respect to marine environment, eradication cases are rare, regarding most occasions from local invasions (see Willan et al. 2000; Wotton et al. 2004; Anderson 2005; Genovesi 2005). In established populations, it is more likely to be expected as a good result the suppression directed at decrease densities rates, which can hold the native ecosystem resilience (Green et al. 2014; Usseglio et al. 2017). The connectivity of aquatic environment and larger dynamic conditions, as currents and depth, complicates the full removal IAS, which are dispersed by many anthropogenic vectors (Bax et al. 2003; Johnston et al. 2009; Ojaveer et al. 2015). This includes those organisms that perform active swimming through water motion, overcoming physicochemical barriers by their permeability and invading new places, even distant (e.g. oceanic islands) or on isolated biogeographical provinces (Floeter et al. 2008; Johnston et al. 2009).\u003c/p\u003e \u003cp\u003eManagement strategies against IAS can proceed in many ways and each approach must consider the interlocution through global and regional protocols, assessing the IAS features to tailor all efforts for successful reduction (Thresher and Kuris 2004; Giakoumi et al. 2019). To note different aspects (e.g. dispersal capacity, movement ecology, distribution in the non-native area) and the level of invasion, managers can adopt different strategies and prioritize resources and technical applications (Hulme 2006). From there, the main groups involved in the IAS problem can adopt well-founded and scientifically supported actions, with the selection of financial and stakeholder networks. For example, lionfish (\u003cem\u003ePterois\u003c/em\u003e spp.) in the invaded Atlantic areas are usually managed through physical removals with the participation of local divers and fishers (Green et al. 2014; Usseglio et al. 2017). Public participation contributes to the involvement of society, as is the case with recreational divers engaged in citizen-science projects, carrying out biodiversity surveys, monitoring the health of reef environments, and managing IAS (Pocock et al. 2014; Thiel et al. 2014; Branchini et al. 2015; Pasternak et al. 2019; Earp and Liconti 2020; Hermoso et al. 2021). This way of controlling IAS combines standardized protocols with regional planning, which is considered an effective method for broadening management efforts (e.g. Frazer et al. 2012; de Le\u0026oacute;n et al. 2013).\u003c/p\u003e \u003cp\u003eNative to the Indo-Pacific, the lionfish \u003cem\u003ePterois volitans\u003c/em\u003e and \u003cem\u003eP. miles\u003c/em\u003e are well-documented species with their invasion tracked over the last few decades, notoriously through north-southwestern Atlantic Ocean expansion in the USA, through the Caribbean region, and more recently on the Brazilian marine waters (Schofield 2010; Johnston and Purkis 2011; C\u0026ocirc;t\u0026eacute; et al. 2013; Ferreira et al. 2015; Luiz Jr. et al. 2013, 2021; Soares et al. 2023; 2025). It is a typical R-strategy case with elevated reproduction rates, a highly dispersive larval stage, and a generalist predator, with wide occurrence in different environments (e.g. reefs, seagrass, estuaries, mesophotic and anthropic reefs) and conditions (e.g. temperature, depth, salinity) (Albins and Hixon 2008; Johnston and Purkis 2011; Claydon et al. 2012; Jud and Layman 2012; Layman and Allgeier 2012; Jud et al. 2015). Population establishment in non-native areas is notably higher, with density rates exceeding native areas several times over (Darling et al. 2011; Kulbicki et al. 2012; Sandel et al. 2015). So, reducing local abundance of lionfish for minimizing the negative impact on non-native areas is the focus of many institutions, and an efficient way of achieving this goal is to involve environmental agencies with the local community in a participatory way. However, most of the data and information on effective management actions and population-reduction measures for lionfish come from the Caribbean, where the invasion has been underway for about 30 years (Claydon et al. 2012; Frazer et al. 2012; Albins and Hixon 2013; de Le\u0026oacute;n et al. 2013; Usseglio et al. 2017).\u003c/p\u003e \u003cp\u003eSince 2020, several lionfish (\u003cem\u003ePterois\u003c/em\u003e spp.) sightings across the Southwestern Atlantic coast (~\u0026thinsp;4,000km) revealed to have successfully established population, including at least 18 Marine Protected Areas (MPAs) (Soares et al. 2023; 2025). The Chico Mendes Institute of Biodiversity Conservation (ICMBio), a Brazilian government agency and part of the Ministry of the Environment and Climate Change, has as its main function the administration of protected natural areas, such as important MPAs. In addition, the team manages the list of threatened species, implements action plans at national level and deals with the context of IAS and the impacts they cause (ICMBio 2023). Considering the history of invasions in different regions of the country, there has been institutional progress regarding the IAS issue and the development of a national policy, which has made it possible to guide management work across the country (Brasil 2019). More specifically, the Fernando de Noronha Archipelago is part of the MPAs managed by the N\u0026uacute;cleo de Gest\u0026atilde;o Integrada de Fernando de Noronha (ICMBio-Noronha), and by state and regional institutions that deal with territorial use of the urban area. With the arrival of the lionfish in Brazil (Ferreira et al. 2015), it was possible to implement various long-term actions for the management of this IAS with a focus on Fernando de Noronha, including prevention, containment, and mitigation to control the population. Since the first record (Luiz Jr. et al. 2021), the lionfish invasion has been monitored locally with the cooperation of several stakeholders, such as Dive Centers, non-governmental institutes, universities, and local community.\u003c/p\u003e \u003cp\u003eThus, this novel research contributes as a considerable example of IAS management in the South Atlantic. Here we discuss the context of the lionfish invasion in the Fernando de Noronha Archipelago, including how we applied the management strategy to mitigate it. We aimed to 1) report on the history of the strategy applied to lionfish management and 2) describe the temporal and spatial variation recorded by multi-staff agents as a result of the presence of this new population in an isolated South Atlantic archipelago. In this context, this is the first article to discuss long-term (2020\u0026ndash;2025) information on lionfish (\u003cem\u003ePterois volitans\u003c/em\u003e) management in the South Atlantic, providing new data on ecological drivers (e.g. currents, depth, substrate) on occurrence, lionfish population density and size.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cp\u003eStudy area\u003c/p\u003e \u003cp\u003eFernando de Noronha (c.03\u0026deg;52\u0026prime;S; 32\u0026deg;25\u0026prime;W) is a Brazilian archipelago located 345 km northeast of the state of Rio Grande do Norte, on the mainland, and about 145 km from the nearest territory, Atol das Rocas (c.03\u0026deg;15\u0026rsquo;S, 33\u0026deg;40\u0026rsquo;W). It is the largest oceanic island in Brazil with 21 islands (26 km\u0026sup2; of total area; 17.6 km long and 60 km perimeter), raised to a depth of 4,000 m and an altitude of 323 m from its volcanic origin in the Fernando de Noronha Submerged Chain, a group of mountains aligned with the continental shelf (Rocha 1995; Linsker 2003; Teixeira et al. 2003). Classified as an Awi Koppen tropical climate, the region is also under the influence of easterly surface winds that affect conditions mainly on windward cost to the south-east, in addition to swell conditions from leeward coast to the north-west between December to March (Eston et al. 1986; Gouveia et al. 2009; Mello et al. 2023). As for the marine environment, the reef consists of a consolidated substrate (i.e. rocky reef) up to 30 m, covered predominantly by turf, coralline, and macroalgae (Eston et al. 1986; Krajewski and Floeter 2011; Matheus et al. 2019). The deeper zones have a mostly sandy composition with the presence of sponges and rhodolith beds, as the coral cover increases to ~\u0026thinsp;5\u0026ndash;20% along a depth gradient (Amado-Filho et al. 2012; Matheus et al. 2019). Fernando de Noronha has around 250 recorded fish species, including endemics and threatened ones (Floeter et al. 2008; Pinheiro et al. 2018; Pimentel et al. 2020), such Rocas gregory \u003cem\u003eStegastes rocasensis\u003c/em\u003e, Noronha wrasse \u003cem\u003eThalassoma noronhanum\u003c/em\u003e and the parrotfish \u003cem\u003eSparisoma\u003c/em\u003e sp., genres already verified in the diet composition items of the lionfish in non-native areas (Eddy et al. 2016; Savva et al. 2020; Murillo-P\u0026eacute;rez et al. 2021; del R\u0026iacute;o et al. 2022) and also in stomach context analysis from lionfish from Fernando de Noronha Archipelago, Brazil (Xavier et al. pers. com.).\u003c/p\u003e \u003cp\u003eIt is the only oceanic island in Brazil with permanent human community, with increasing c. 4,000 people (IBGE 2023), and the Protected Areas in the archipelago's territory include two distinct uses: the Environmental Protected Area (APA; sustainable use; MMA, 2017), and the National Marine Park of Fernando de Noronha (PARNA; strict protection; IBAMA 1990). This last demarks the delimitation over the 50-meter isobath in the marine area. Recreational SCUBA divers visit the two protected areas at different sites, most at depths of up to 20 m, and eventually use open circuits, rebreathers or technical diving for access deeper and longer dives. On-board operations are conducted by Diving Centers (DC), three of which are authorized for commercial guided tourism in the PARNA (numbered as DC1\u0026ndash;3), in addition to one focused on lionfish management in this same area and APA (DC4). Five Image Companies (IC) are authorized to accompany DC operations, focusing on selling photo and video media. Diving operations are organized usually with two dives per shift and company (morning and afternoon, with night dives less frequently) with c.50 minutes per site. This results in an average of ten boat operations and 20 active groups of divers per day in the Archipelago, reaching c.10,000 dives per year. Coastal SCUBA diving is also allowed daily in the port region, and freediving is one of the commonest tourism activities in shallow areas close to the main island.\u003c/p\u003e \u003cp\u003eFernando de Noronha has a history involving terrestrial IAS (see Micheletti et al. 2020), but there have never been reports of IAS in the marine environment. In terms of invasive expansion, the lionfish was introduced in Florida in the 1980s and has been increasing its occupation of the South Atlantic (Schofield 2010; Johnston and Purkis 2011). In December of 2020, the species arrived in the Fernando de Noronha Archipelago (Luiz Jr. et al. 2021) and is moving on to other coastal areas (Soares et al. 2022; Maggioni et al. 2023).\u003c/p\u003e \u003cp\u003eStrategic alignment according to legal terms\u003c/p\u003e \u003cp\u003eIn dealing with a new IAS invasion, we report on the application of the management strategy, including all steps for the arrival of the lionfish in Fernando de Noronha archipelago. Preventive works were carried out before the invasion by implementing educational initiatives in the Fernando de Noronha archipelago. After the first sighting of lionfish on the island, we follow the proceedings based on Ordinance No. 19/2025 (Brazil, 2025), a legal instrument that regulates nationally the prevention and management actions for the control and eradication of IAS in Natural Protected Areas and their surroundings. The standardized technical document is officialized as the \u0026ldquo;Management Project\u0026rdquo; (MP), which was carried out by the ICMBio-Noronha to its validation by higher authorities. This includes standard actions, dissemination and education activities, specimen destination, adequate training and license for the capture of individuals, and the permission of local agents not directly linked to the local environmental agency was able to participate spontaneously, via a consent term (\u0026ldquo;Adhesion Term\u0026rdquo;), to promote the lionfish management. By the collaborative effort of multiple institutions, we call as \u0026ldquo;multi-staff\u0026rdquo; all the stakeholders and team members who contributed to the actions of the management process.\u003c/p\u003e \u003cp\u003eControl based on multi-staff integration\u003c/p\u003e \u003cp\u003eBy consolidating the management strategies, local agents, including DCs, ICs and autonomous (e.g. visitors guide, fishers and residents) staff, were training in the characteristics and methods for individual captures and inscribed to follow specific rules regarding lionfish management. Especially when dive operations was targeting a higher effort to lionfish captures, the following topics were clarified: 1. Diving with a focus on lionfish detection (e.g. lionfish-focused search; Green, 2012); 2. Keeping the dive profile suitable for the methodology, ensuring a proper scanning of the substrate; 3. Carefully observing caves, holes and places with less light conducting right to not impact substrate (i.e. Krieger et al. 2013; Giglio et al. 2020); 4. Paying attention to safety regulations and the risk of accidents involving lionfish; and 5. In the event of a sighting, the person in charge would give an audible or visual signal so that the group could record images and follow with the lionfish capture.\u003c/p\u003e \u003cp\u003eLionfish has a low displacement rate, but they can change location in a short period of time (Jud et al. 2015; Bacheler et al. 2015; Tamburello and C\u0026ocirc;t\u0026eacute; 2015). Thus, we instructed all local agents to take photos of individuals and reference them as best they could. Only properly trained and authorized divers captured the lionfish individuals with appropriate equipment: a 90 cm Hawaiian Sling harpoon with three to five barbs and the containment PVC tube (standard size of c.15 cm diameter x c.60 cm length) for safely storing captured lionfish. The kit stayed on the boat during all diving operations or taken together for freediving occasions, besides thermoses with hot water or a heat pack on field trips, in case of contact with the lionfish\u0026rsquo;s poisonous spines. Therefore, divers reported biotic and abiotic parameters based on the dive computer and personal perception, sending each record through an online form (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). All information and specimens were forwarded to the ICMBio-Noronha team for standardization in data input. Thus, we collected biometrics of weight (g) and total length (cm), identifying individuals with numbered plastic tags or graphite on paper records to froze and later send them to other institutions that undertake specific research (e.g. diet and genetics).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eStandard information for local agents to fill in when recording invasive lionfish, sectioned into sightings and captures, followed by a characterization of the site and biotic and abiotic factors related to recorded individuals in the Fernando de Noronha Archipelago (South Atlantic, Brazil).\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"2\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eN\u0026ordm;\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eInformation\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSighting date:\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSighting time:\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eWho sighted (first and last name):\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHow many individuals were sighted:\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCapture date:\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCapture time:\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eWho captured it (first and last name):\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHow many individuals were captured:\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEquipment used (harpoon or another accessory):\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eInstitute/Diving Center:\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDive site/reference:\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDepth:\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eWater temperature:\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCurrent (weak, moderate, or strong):\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSubstrate (sand, algae, corals, or artificial):\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFish behavior (rest, or swin):\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEstimated size (\u0026lt;\u0026thinsp;10 cm, between 10 and 20 cm or \u0026gt;\u0026thinsp;20 cm):\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eOther details:\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003ePost-fieldwork processing for data analysis\u003c/p\u003e \u003cp\u003eLionfish records were treated as a) captures, being the removal of the individual from the environment using some support instrument; b) sightings, being field observations without capture; and c) suspicions, being inaccurate sightings of lionfish by amateur audiences with no proven identification experience. We do not rule out registering suspicions because this was a preventive and management emergency action. We standardized two operation types of targets to the control of the local invasion: 1) opportunistic records during daily activities, as commercial tourist SCUBA diving, fisheries and recreational visitation (hereafter \u0026ldquo;Routine\u0026rdquo;) and 2) high effort (i.e. Underwater Visual Transects) for lionfish records during SCUBA or freediving (hereafter \u0026ldquo;Monitoring\u0026rdquo;). Group size and time during dives varies widely and was not a predictor between ratings. Therefore, we standardized all localities with lionfish records with a total area of 1.5 hectares (ha; c. 15,000 m\u0026sup2;; adapted from Luiz Jr., 2009) to evaluate lionfish density per point based on recorded events over time. Points visited in the same month were corrected as a single recorded event, to avoid underestimating lionfish density. We trademarked 20 m for categorizing all recorded events as shallow (\u0026lt;\u0026thinsp;20 m) and deep (\u0026gt;\u0026thinsp;20 m) waters, by considering the profile usually performed in dive points by local agents and limits of diving certifications.\u003c/p\u003e \u003cp\u003ePopulation estimate values were evaluated including the total number of records during the study period, by applying the intrinsic growth rate (\u003cem\u003er\u003c/em\u003e): \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:r=\\text{l}\\text{n}\\left[\\left({N}_{t2}/{N}_{t1}\\right)\\right]/(t2-t1)\\)\u003c/span\u003e\u003c/span\u003e, and increment rate (\u003cem\u003eλ\u003c/em\u003e): \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\lambda\\:={e}^{r}\\)\u003c/span\u003e\u003c/span\u003e(e.g., Margalef 1998), where \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\text{l}\\text{n}\\)\u003c/span\u003e\u003c/span\u003e is the logarithm of the fraction between the final (\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:{N}_{t2}\\)\u003c/span\u003e\u003c/span\u003e) and initial (\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:{N}_{t1}\\)\u003c/span\u003e\u003c/span\u003e) population size, and \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:e\\)\u003c/span\u003e\u003c/span\u003e is the base of the natural logarithm (~\u0026thinsp;2.718). We analyzed the length-weight (L-W) relationship using the allometric growth function \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:W=a\\times\\:{\\:L}^{\\:b}\\)\u003c/span\u003e\u003c/span\u003e and later the body condition factor (Fulton\u0026rsquo;s K) through \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:K=(W/{L}^{3})\\times\\:100\\)\u003c/span\u003e\u003c/span\u003e. To better characterize the lionfish detection scenario in the archipelago region, we compared the frequency of records and their respective information. We use multiple regression to check the relation between density by time and depth, factorial ANOVA to relate records between the operation types (routine/monitoring) and depth, as the same for density per month to check inter-annual variation. To compare the complementary data provided by the records, we use Chi-Square with expected proportion tests with no difference between categories, including divers' perceptions about the substrate (benthic organism or soil backgrounded with the fish), marine current (drift speed at each dive point) and fish behavior (individual activity status). Generalized Linear Model (GLM) was applied to compare protected areas, sea regions and body conditions on depth and time.\u003c/p\u003e \u003cp\u003eDetected sites with more than 3 lionfish recorded events were evaluated to compare the effect of management efforts on density by time, using the slope coefficient of linear regression to classify decrease (\u0026lt;\u0026thinsp;0), stable (\u0026asymp;\u0026thinsp;0), and increase (\u0026gt;\u0026thinsp;0) trends. To spatially analyze of lionfish distribution in the Fernando de Noronha Archipelago we estimate the most explored areas for each bathymetry features (20, 50, 100 e 500 m depth) categorizing five classes (0\u0026ndash;70 m as well explored, 71\u0026ndash;100 few explored, 101\u0026ndash;150 inferred, 151\u0026ndash;300 distant, and 301\u0026ndash;4000 very distant) by referring a Euclidian Distance raster. In order to predict the size of the fish population in this area, we considered the maximum density values for each location with a recorded event and used the geostatistical interpolation method by ordinary kriging. The values were transformed using the natural logarithm (ln) for better adjustment of the semivariogram Gaussian model. Subsequently, the data was reprocessed for spatial analysis of the species density distribution. The population estimate was based on the sum of the estimated density values in all cells (100 m\u0026sup2;) within the study area. We used QGis v3.40 ('r.grow.distance' and 'Smart-Map' tools) for the spatial analyses and map output and RStudio v.2023.04.1 for the statistical tests and visual graphics (α\u0026thinsp;=\u0026thinsp;0.05).\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eReporting on management strategy processes\u003c/p\u003e \u003cp\u003eWe characterized five main steps based on Brazilian policy scenario for IAS management protocol (Ordinance No. 19/2025), including the period before the lionfish invasion begins to the stabilization of its management strategy: 1) prevention about IAS; 2) IAS record; 3) preparation and submission of MP; 4) evaluation and MP approval; and 5) upon approval, implementation of the \u0026ldquo;Adhesion Term\u0026rdquo; in order to regulate the management actions by local agents (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). All the actions for the lionfish in Fernando de Noronha were taken by the engagement of multiple institutions to procedure with its management (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBefore the arrival of the lionfish in the Fernando de Noronha Archipelago, previous publications focused on prevention for Brazil (e.g. ICMBio 2019) and relevant actions were also made locally in 2018\u0026ndash;2019 by Projeto Conserva\u0026ccedil;\u0026atilde;o Recifal (PCR; \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://conservacaorecifal.com/\u003c/span\u003e\u003cspan address=\"https://conservacaorecifal.com/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), focusing on educational actions and training the government agencies, divers, fishermen, residents, and tourists, about the lionfish threat as IAS (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA). The first lionfish was detected on 20 December 2020, supported by the Universidade Federal Fluminense (UFF) with DC4 (under authorization SISBIO 41327; Luiz Jr. et al. 2021). To enabled rapid communication and first actions with local agents (mainly the DCs), the \u0026ldquo;Emergency strategy for lionfish management in the Fernando de Noronha Archipelago\u0026rdquo; (ICMBio 2021) was consolidated by the ICMBio-Noronha on 30 March 2021. This strategic protocol had resulted in the initial training and lionfish captures (n\u0026thinsp;=\u0026thinsp;25), while the MP was in progress. The preparation and submission of MP was made by ICMBio-Noronha on 6 October 2021, which was ratified 30 days after submission. This made up the ongoing instructional procedure that regulates the multi-staff agents (108 members and all the four DC which works onboard) to act via \u0026ldquo;Adhesion Term\u0026rdquo; on lionfish management in Fernando de Noronha. Since this period, it was totalized 2,660 lionfish records in all Archipelago. In addition, on 10 September 2024, Ordinance No. 2,761 revised the Management Plan of the Fernando de Noronha Marine National Park (i.e. IBAMA 1990). This regulatory amendment enabled IAS control activities associated with tourism-related operations throughout legally and autonomously participate of dive operations, providing a complementary mechanism to support the financial sustainability of ongoing control efforts.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eProcess for structuring an IAS Management Project (MP), according to the Brazilian policy scenario (Ordinance No. 6/2019), compared with the lionfish invasion in the Fernando de Noronha Archipelago (this paper) and the management effort of fieldwork over time.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eStep 1\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eStep 2\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eStep 3\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eStep 4\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eStep 5\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eTotal\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eManagement process based on Brazilian guidelines\u003c/b\u003e\u003csup\u003e\u003cb\u003e*\u003c/b\u003e\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePrevention about IAS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIAS record\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePreparation and submission of MP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEvaluation and MP approval\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAdhesion Term for local agents\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eTotal\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eLionfish case in Fernando de Noronha\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2018\u0026ndash;2019\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e20 December 2020\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6 October\u003c/p\u003e \u003cp\u003e2021\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5 November 2021\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e18 November 2021 (ongoing)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e20 December 2020 to 30 September 2025 (ongoing)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eDuration of the process\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e291 days\u003c/p\u003e \u003cp\u003e(~\u0026thinsp;11 months)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e321 days\u003c/p\u003e \u003cp\u003e(~\u0026thinsp;12 months)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e334 days\u003c/p\u003e \u003cp\u003e(~\u0026thinsp;12 months)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1,746 days\u003c/p\u003e \u003cp\u003e(~\u0026thinsp;58 months)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eNumber of lionfish records\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2,660\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eValidated document for management\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEmergency strategy\u003c/p\u003e \u003cp\u003e(ICMBio 2021)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eEmergency strategy\u003c/p\u003e \u003cp\u003e(ICMBio 2021)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEmergency strategy\u003c/p\u003e \u003cp\u003e(ICMBio 2021)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eManagement Project\u003c/p\u003e \u003cp\u003e(ongoing)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eSupport lionfish management was performed between the multi-staff institutions covered by specific roles for decision-making from local to international level (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Permission and institutional orientation through MP alignment were provided on a national scale by the high instance of national government represented by Coordination of Invasive Alien Species Management (ICMBio-CMEEI). From a regional scale, technical guidance, planning and specific research (including specimen destination) was subsided by Non-governmental organization PCR and Universities, such as Universidade Federal Fluminense (UFF), Universidade Federal do Rio Grande de Norte (UFRN), Universidade Federal de Pernambuco (UFPE), Universidade Federal de Alagoas (UFAL), and Universidade Federal do Cear\u0026aacute; (UFC), besides network engagement for planning actions by Pernambuco State Environment Agency (SEMAS/CPRH; Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eC). International support included key capacity building through inter-institutional exchanges with experts (represented by PB) of Bonaire\u0026rsquo;s Environmental Agency, the Stichting Nationale Parken Bonaire (STINAPA; Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB), as well as specific research of California Academy of Sciences (CAS). Thus, the ICMBio-Noronha assumed a central role to coordinate multiple institutions, overall, to maintain regular training sessions, processing data and specimens and stimulate lionfish management with local DCs and ICs, which provide most lionfish records during routine and monitoring operations (n\u0026thinsp;=\u0026thinsp;2,614; 98.3%; Fig. D\u0026ndash;F).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eResults outcome by multi-staff integration\u003c/p\u003e \u003cp\u003eDuring the period from 20 December 2020 to 30 September 2025, there were registered 2,404 captures, 251 sightings and 5 suspicions (N\u0026thinsp;=\u0026thinsp;2,660) of lionfish in the Fernando de Noronha Archipelago, featuring an exponential increase of 345.9% in its population over this period (\u003cem\u003er\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1.495; \u003cem\u003eλ\u003c/em\u003e\u0026thinsp;=\u0026thinsp;4.459; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eA). It was observed that the highest value reached 299 records in September 2025 (292 captures), followed by June 2025 (n\u0026thinsp;=\u0026thinsp;213; 208captures) and April 2025 (n\u0026thinsp;=\u0026thinsp;126; 125captures). After the first record in December 2020, there were no detections for six months of lionfish individuals, when the second individual was captured in June 2021, presenting a continuous increase every month since then (45.1\u0026thinsp;\u0026plusmn;\u0026thinsp;56.3 records/month). Through 52 months, records amounted to 407 events (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eB), gradually increasing each year until they peak in 2025 between shallow (n\u0026thinsp;=\u0026thinsp;184 records; 44 events) and deep waters (n\u0026thinsp;=\u0026thinsp;1030 records; 85 events). Regarding inter-annual variations, lionfish densities have not varied significantly between months (F\u003csub\u003e11,73\u003c/sub\u003e = 0.526; p\u0026thinsp;=\u0026thinsp;0.874). Moreover, we could check that densities were related positively with time and deeper records (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.521; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001), by presenting higher densities in deep waters (8.2\u0026thinsp;\u0026plusmn;\u0026thinsp;3.9 individuals/ha) than compared with shallow regions (2.5\u0026thinsp;\u0026plusmn;\u0026thinsp;2.1 individuals/ha) in recent years (t\u003csub\u003e2025\u003c/sub\u003e). Thus, the general scenario of lionfish invasion is estimated at 3.9 individuals/ha for the Fernando de Noronha Archipelago.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eDepth gradient showed a normal distribution (33.0\u0026thinsp;\u0026plusmn;\u0026thinsp;12.1 m), counting with more individuals between 20 to 40 m (n\u0026thinsp;=\u0026thinsp;1,593; 62.3%) and less records in shallower (34 records above 10 m; min.: 0.5 m) and deeper (130 records under 50 m; max.: 95 m) areas. Records were related significantly with deep waters (F\u003csub\u003e2,1726\u003c/sub\u003e = 16.010; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and monitoring operations (F\u003csub\u003e1,4937\u003c/sub\u003e = 45.795; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001), which presented major values and variation for the number of records (6.9\u0026thinsp;\u0026plusmn;\u0026thinsp;9.3 ind./ha) than opportunistic routine records (1.8\u0026thinsp;\u0026plusmn;\u0026thinsp;2.1 ind./ha). In addition, SCUBA diving was the most collaborative activity for field detection in all Archipelago (n\u0026thinsp;=\u0026thinsp;2,634; 99%), with few records made up during freediving (n\u0026thinsp;=\u0026thinsp;24; 0.9%). A single catch by a fisherman was verified and one case when the lionfish was filmed almost at the surface by a person above water at the Port (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Thus, recorded lionfish were mainly performed by Dive Instructors (n\u0026thinsp;=\u0026thinsp;2,597; 97.6%), and, less representative, by other agents, such as camera divers, tourists, fishers, and residents (n\u0026thinsp;=\u0026thinsp;63; 2.3%). Great effort was realized by DC4 (n\u0026thinsp;=\u0026thinsp;2,278; 85.6%), mainly related to focused operations on monitoring deep areas in the Archipelago, followed by those carrying out largely routine operations by DC1\u0026ndash;3 and ICs (n\u0026thinsp;=\u0026thinsp;336; 12.6%). Although not always provided, complemented information was noted several occasions by multi-staff registers, regarding most records for I) recording time (n\u003csub\u003e[morning]\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;584; 53.4%), II) marine current (n\u003csub\u003e[weak]\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;1911; 91.3%), III) substrate composition (n\u003csub\u003e[coral]\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;1,012; 49.6%), and IV) fish behavior (n\u003csub\u003e[rest]\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;1963; 99.4%).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eData parameters on lionfish records in the Fernando de Noronha Archipelago gathered by local agents, including the operation type, related activity, local agents, Dive Centers (DC), recording time, marine current, substrate composition, fish behavior, and Protected Natural Areas, by categories of suspect, sighting, and captures. Each section is accompanied by the Chi-Square result and its significance level (* represents the significant category).\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCategories\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSuspect\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSighting\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCaptures\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eTotal\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c5\" namest=\"c1\"\u003e \u003cp\u003e\u003cb\u003eMethod\u003c/b\u003e (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;1295.0; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001; N\u0026thinsp;=\u0026thinsp;2,660)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMonitoring *\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e105\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2,153\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2,258 (84.9%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRoutine\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e146\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e251\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e402 (15.1%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c5\" namest=\"c1\"\u003e \u003cp\u003e\u003cb\u003eActivity\u003c/b\u003e (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;7773.9; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001; N\u0026thinsp;=\u0026thinsp;2,660)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSCUBA diving *\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e242\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2,392\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2,634 (99%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFreediving\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e24 (0.9%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFishing\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1 (0,04%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAbove water\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1 (0.04%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c5\" namest=\"c1\"\u003e \u003cp\u003e\u003cb\u003eLocal agents\u003c/b\u003e (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;10021.0; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001; N\u0026thinsp;=\u0026thinsp;2,660)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDive Instructors *\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e216\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2,381\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2,597 (97.6%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCamera divers\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e19 (0.7%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTourists\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e6 (0.2%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFishers\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1 (0.04%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOthers\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e37 (1.4%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c5\" namest=\"c1\"\u003e \u003cp\u003e\u003cb\u003eDive Centers\u003c/b\u003e (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;9142.6; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001; N\u0026thinsp;=\u0026thinsp;2,660)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDC1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e75 (2.8%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDC2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e130\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e171 (6.4%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDC3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e78 (2.9%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDC4 *\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e117\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2,161\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2,278 (85.6%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eICs\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e12 (0.5%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOthers\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e46 (1.7%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c5\" namest=\"c1\"\u003e \u003cp\u003e\u003cb\u003eRecording time\u003c/b\u003e (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;6.8; p\u0026thinsp;=\u0026thinsp;0.009; N\u0026thinsp;=\u0026thinsp;1,086)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMorning *\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e84\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e499\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e586 (54.0%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAfternoon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e462\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e500 (46.0%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c5\" namest=\"c1\"\u003e \u003cp\u003e\u003cb\u003eMarine current\u003c/b\u003e (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;3186.3; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001; N\u0026thinsp;=\u0026thinsp;2,099)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWeak *\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e156\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1,754\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1,915 (91.2%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eModerate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e158\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e175 (8.3%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eStrong\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e9 (0.4%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c5\" namest=\"c1\"\u003e \u003cp\u003e\u003cb\u003eSubstrate composition\u003c/b\u003e (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;1964.7; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001; N\u0026thinsp;=\u0026thinsp;2,039)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSand\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e80 (3.0%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAlgae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e446\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e474 (17.8%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCoral *\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e85\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e927\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1,012 (38.1%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRock\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e242\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e290 (10.9%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArtificial\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e44 (1.7%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePorifera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e133\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e139 (5.2%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c5\" namest=\"c1\"\u003e \u003cp\u003e\u003cb\u003eFish behavior\u003c/b\u003e (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;1927.3; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001; N\u0026thinsp;=\u0026thinsp;1,975)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRest *\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e162\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1,797\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1,963 (73.8%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSwin\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e12 (0.5%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c5\" namest=\"c1\"\u003e \u003cp\u003e\u003cb\u003eProtected Areas\u003c/b\u003e (χ\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;320.9; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001; N\u0026thinsp;=\u0026thinsp;2,660)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAPA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e73\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e794\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e868 (32.6%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePARNA *\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e178\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1,610\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1,792 (67.4%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eTotal\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e5 (0.2%)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e251 (9.4%)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e2,404 (90.4%)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e2,660 (100%)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eIndividuals generally average 23.17\u0026thinsp;\u0026plusmn;\u0026thinsp;6.7 cm of total length, recording smaller (8.1 cm) to bigger (46.0 cm) detections. Weight remained between 238.2\u0026thinsp;\u0026plusmn;\u0026thinsp;241.6 g, ranging from 7.0 to 1670.0 g. There was a significant difference over the years for both parameters, with higher values for size (F\u003csub\u003e5, 1275\u003c/sub\u003e = 18.594; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and weight (F\u003csub\u003e5, 1275\u003c/sub\u003e = 10.986; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001) in recent years. The L-W relationship showed a strong association (R\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;0.96; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and high coefficient (b\u0026thinsp;=\u0026thinsp;3.22), pointing to positive allometric growth and larger individuals presenting proportionally more body mass in relation to length (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eA). The body condition factor reveals high physiological condition (Fulton\u0026rsquo;s K\u0026thinsp;=\u0026thinsp;1.43). The effect of time on the condition factor K was significant (F\u003csub\u003e5, 1275\u003c/sub\u003e = 16.76, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001), indicating that the body condition of the population varied between years (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eB). Despite the wide variation, body size was significantly related to depth, with higher averages (K: 1.38\u003csub\u003e[deep]\u003c/sub\u003e\u0026thinsp;\u0026gt;\u0026thinsp;1.31\u003csub\u003e[shallow]\u003c/sub\u003e) for individuals at greater depths (F\u003csub\u003e1, 606\u003c/sub\u003e = 7.028; p\u0026thinsp;=\u0026thinsp;0.008).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eMult-staff effort totalized 116 localities, which represented c.610.50 ha throughout the 407 recorded events during this study period. Lionfish individuals were recorded in both protected areas of PARNA (n\u0026thinsp;=\u0026thinsp;1,792; 67.4%) and APA (n\u0026thinsp;=\u0026thinsp;868; 32.6%), besides regions of Leeward (n\u0026thinsp;=\u0026thinsp;2,289; 86.2%) and Windward (n\u0026thinsp;=\u0026thinsp;367; 13.8%) Sea regions. Species density was significantly greater in the PARNA (5.2\u0026thinsp;\u0026plusmn;\u0026thinsp;8.7 ind./ha) than in the APA (3.7\u0026thinsp;\u0026plusmn;\u0026thinsp;4.9 ind./ha) areas (β\u0026thinsp;=\u0026thinsp;0.338; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001), and for leeward (5.0\u0026thinsp;\u0026plusmn;\u0026thinsp;8.2 ind./ha) than in windward (2.9\u0026thinsp;\u0026plusmn;\u0026thinsp;3.1 ind./ha) sea regions (β = \u0026minus;\u0026thinsp;0.522; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Most of recorded events showed lower densities (n\u0026thinsp;=\u0026thinsp;305; 78.6%), ranging from 1 to 5 individuals per point. Higher densities were found in three unusual points located in Leeward Sea and in the PARNA, counting respectively with 90 (60 ind./ha), 80 (53 ind./ha), and 63 (42 ind./ha) lionfish records in single visits to the dive sites. Official dive sites (i.e. authorized for commercial visitation) were surveyed in 18 localities and represent 349 lionfish records. These points presented lower densities in general (2.1\u0026thinsp;\u0026plusmn;\u0026thinsp;3.3 ind./ha), although high detections were checked in Cagarras Fundas (n\u0026thinsp;=\u0026thinsp;101 records), Laje Dois Irm\u0026atilde;os (n\u0026thinsp;=\u0026thinsp;81), and Cabe\u0026ccedil;o da Sapata (n\u0026thinsp;=\u0026thinsp;59) over the study period. The effect of field management proved to be effective in maintaining a low temporal trend (slope coefficient median\u0026thinsp;=\u0026thinsp;0.02) throughout the evaluated period (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Most of repeatedly visited sites (n\u003csub\u003e[S1\u0026ndash;S45]\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;45; 6.51\u0026thinsp;\u0026plusmn;\u0026thinsp;4.70 visits/site) showed stable trends (n\u0026thinsp;=\u0026thinsp;30; \u003cem\u003ea\u003c/em\u003e\u0026thinsp;\u0026asymp;\u0026thinsp;0) with low density averages (2.01\u0026thinsp;\u0026plusmn;\u0026thinsp;1.68 ind./ha), although there has been an overall average increase (0.10\u0026thinsp;\u0026plusmn;\u0026thinsp;0.46 coefficient units) and some locations (n\u0026thinsp;=\u0026thinsp;10; 22%) recorded pronounced growth (\u003cem\u003ea\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;0.13).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eEuclidean distance map presented a general value of 1,364\u0026thinsp;\u0026plusmn;\u0026thinsp;956 m (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e), pointing to 17,331.98 ha (89.4%) of entire study area (0\u0026ndash;500 m depth) as very distance regions (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Well explored areas (181.51 ha; 0.9%) was checked over 20 m (98.91 ha; 54.1%), followed by 21 m to 50 m (68.82 ha; 37.9.0%), 51 m to 100 m (10.87 ha; 6.0%), and 500 m to 100 m (2.91 ha; 1.6%), besides few explored areas (172.13 ha; 0.9%), which totalizing both 353.64 ha (1.8%) of most recognized areas surround the Archipelago. The adjusted variogram model showed a good predictive capacity (RMSE\u0026thinsp;=\u0026thinsp;0.414; R\u0026sup2; = 0.661) and positive spatial autocorrelation by presenting local density clusters through interpolation analysis (Moran's I\u0026thinsp;=\u0026thinsp;0.161; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). The kriging map showed the presence of areas of higher density (above 20 ind./ha) concentrated in specific sectors of leeward sea, with a decreasing gradient toward the remote areas (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e). Thus, the population estimate using this analysis reached 75,742 (CI 95% 56,202\u0026ndash;95,372) individuals and lionfish density of 3.90 (CI 95% 2.89\u0026ndash;4.91) ind./ha in Fernando de Noronha Archipelago.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eEstimated lionfish sampling areas (ha; percentage) into five effort classes (well explored: 0\u0026ndash;70 m; few explored: 71\u0026ndash;100 m; inferred: 101\u0026ndash;150 m; distant: 151\u0026ndash;300 m; very distant: 301\u0026ndash;4000 m) and between the sectors of the isobath lines from 0 to 20, 21 to 250, 51 to 100, and 101 to 500 m depth.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eEffort classes\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"4\" nameend=\"c5\" namest=\"c2\"\u003e \u003cp\u003eIsobaths classes (m)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eTotal\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u0026ndash;20\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e21\u0026ndash;50\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e51\u0026ndash;100\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e101\u0026ndash;500\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWell explored\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e98.91 (54.5%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e68.82 (37.9%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10.87 (6.0%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2.91 (1.6%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e181.51 (0.9%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFew explored\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e86.23 (50.1%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e71.97 (41.8%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e11.23 (6.5%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2.7 (1.6%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e172.13 (0.9%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInferred\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e168.86 (45.6%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e167.47 (45.2%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e26.86 (7.3%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e6.96 (1.9%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e370.15 (1.9%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDistant\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e476.34 (36.1%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e661.01 (50.0%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e145.86 (11.0%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e37.52 (2.8%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1,320.73 (6.8%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eVery distant\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1,151.29 (6.6%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e8,047.75 (46.4%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e3,751.19 (21.6%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4,381.75 (25.3%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e17,331.98 (89.4%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eTotal\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e1,981.63 (10.2%)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e9,017.02 (46.5%)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e3,946.01 (20.4%)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e4,431.84 (22.9%)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e19,376.5 (100%)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe management strategy presented herein demonstrates that the use of legal and protocolary adoptions have efficiently stabilized and outlined the lionfish control through standardized actions in the Fernando de Noronha Archipelago. Combining multi-staff actions based on international and national protocols with a strong local field effort, it was possible to designate specific roles covering better decision-making. Consequently, these actions generated positive results for understanding lionfish population and promoting a protocol for local agents, which was reflected in the number of records and practical management in the South Atlantic. Even considering the high population level locally, we could check that the effort in several sites was effective to control lionfish density throughout the time, including a decrease over time in some areas. Those results reflect the main strategy to be maintained in areas affected by lionfish invasion, even as a form of local control, in addition to encouraging management of the individuals captures (Green et al. 2014; Bogdanof et al. 2021). In comparison to other locations, we observed that similar protocols have also been effective for more complex and intense lionfish invasions, such as Caribbean islands and Mediterranean Sea (Dahl et al. 2019; Ulman et al. 2022). We highlight that the management strategy for lionfish control in FNA constitutes a model to be applied in different locations, which must be adapted according to the scale to be followed and its regulations.\u003c/p\u003e \u003cp\u003eLionfish management in Fernando de Noronha showed that SCUBA diving and the active participation of dive instructors were key factors in controlling the species. These results are like other regions that have shown the integration of various teams in lionfish removal actions (Anderson et al. 2017). This involved the entire time series related to the detection of this IAS, as observed since the first record of the species in 2020. All steps of the management project were designed to include divers in effective control, given that this activity is one of the most practiced in the archipelago and that the areas most accessed by citizen scientist divers are shallow coastal subtidal areas, with depths\u0026thinsp;\u0026lt;\u0026thinsp;40 m (Thiel et al. 2014). Freediving participation was also significant, although the number of records was lower. This may be related to the higher density in deep areas compared to shallow waters. Furthermore, there was a significant increase in the number of individuals below 20 m depth, especially in 2025, noting that the density trend in shallow waters can increase more slowly over time. This pattern is recognized for lionfish invasion processes, and their colonization may differ between depth levels, with levels in shallow and deep waters becoming similar over time (Airey et al. 2023).\u003c/p\u003e \u003cp\u003eAlthough we have observed stable trends in several areas, density trends at some sites and the cumulative curve show that lionfish occupation in general is progressing rapidly and is probably greater than management efforts can support. The pattern in IAS begins with low-density introduction and establishment levels and exponentially increasing density and population growth rates (Sakai et al. 2001; Dahl et al. 2019), a similar pattern observed in these early years of lionfish recognition in our study site. In addition, our work focuses mainly on relative effort measures, based on the number of detections provided in each record reported by local agents. It should be noted that some sites receive continuous visits, such as official commercial dive sites and shallower areas frequented by freedivers, which is considered a significant effort. However, by checking nested clusters for the spatial distribution of lionfish population in Fernando de Noronha, we can infer that other biological variables may have a stronger influence on distribution throughout the archipelago. Lionfish populations are denser in deeper waters, related to the range of habitat diversity, main areas of population recruitment due to site fidelity, influence of light penetration, as well as lower pressure from the removal of individuals, with the dispersion of individuals aggregating from deep to shallow waters (Biggs and Olden 2011; Nuttal 2014; Airey et al. 2023). Despite the effort undertaken in this study, there was less frequent diving in mesophotic regions, which presented greater technical difficulty and consequently a lag in the effort to estimate individuals.\u003c/p\u003e \u003cp\u003ePrevious studies have demonstrated that lionfish populations frequently reach higher densities and comprise larger individuals within marine protected areas (MPAs), when compared to adjacent non-protected zones (Morris et al. 2011; de Le\u0026oacute;n et al. 2013; Andradi-Brown et al. 2017; Kleitou et al. 2024). In this regard, lionfish currently occupy 33% of Brazil\u0026rsquo;s MPAs and are likely to spread to an additional 25 protected areas (c.60% of MPAs) within the next decade, while regular removal programs are absent in most of them (Soares et al. 2025). Such findings emphasize the challenge of managing lionfish in protected areas, where no-take zones should be specially treated with IAS removal through management strategies because restrictions on extractive activities create favorable conditions for invasive populations to persist. As observed, lionfish distribution in Fernando de Noronha is not specifically related to the boundaries of protected areas. High and low-density points were found in both visited and unvisited areas, which reinforces that the sampling effort applied for management should be applied more systematically. In this case, the interpretations obtained to affect population control can be better grounded in ecological factors and through planned mapping. This also reinforces decision-making regarding invasion areas, showing that continuous monitoring and local agents training, especially diving instructors, should be done more frequently. Considering the negative impacts caused by the intensity of recreational diving in various locations (Giglio et al. 2020; 2025), caution and good planning are needed to open new sites, especially in protected areas (Soares et al. 2023). We therefore encourage lionfish management actions to be carried out with the support of planned strategies. For instance, considering sea conditions and diving safety, it would be optimal to apply a grid of plots over the marine region of the archipelago and sampling these locations through monitoring operations, by applying focused census and proper removal of lionfish.\u003c/p\u003e \u003cp\u003eThe role of diving operators proved to be highly relevant in ensuring that IAS control was more successful. It was observed that one Dive Center (DC4) made a great effort regarding the number of records, including the lionfish sightings and, mainly, captures. This was related that DC4 maintains continuous operations with a greater focus on lionfish control, being recognized and legally authorized to operate in the protected areas around the archipelago. This strategy was well interpreted in terms of management strategy due to the difference between diving operation methods. Operations in the PARNA is performed historically before lionfish invasion by three specific dive centers (DC1\u0026ndash;3), as this is a region that requires greater control due to its environmental sensitivity. Considering that the recreative commercial model of these operations was focused on customer service and recreational diving safety, we call them routine operations, which also contributed in an important way to the mapping of records in shallow waters and at official sites in PARNA. Thus, it became quite strategic to regulate the format of operations focused on lionfish capture, referred to here as monitoring operations, resulting in greater recognition of sites and increasing the number of records. Similarly, these activities must be well coordinated so as not to cause overlaps between groups and greater impact on diving spots (Luiz Jr. 2009). In addition, the routine operations are mainly carried out in shallow waters, which have been ecologically linked to lower lionfish detection rates. By observing lionfish increase over time at these sites, it is crucial to conduct more operations focused on species control, balancing them with recreational operations.\u003c/p\u003e \u003cp\u003eLocal communication was also a distinctive factor in making IAS control practical. Population control and action protocols tend to be better managed and controlled in places with a smaller territorial scale, due to the possibility of applying management efforts in the field and communication between local agents in a simpler and more effective way (Earp and Liconti 2020). This reflects in better data updating and, thus, in the understanding of parameters related to the lionfish population (Kelly et al. 2020). We consider that the reporting of records via virtual app was effective in proceeding with the analyses applied here, which should be further encouraged, so that the number of registered individuals could be disclosed faster by interested media and all teams involved.\u003c/p\u003e \u003cp\u003eRegarding the management permission process, despite the extensive bureaucracy of Brazilian public processes in general, this procedure was carried out with reasonable promptness. Since the first lionfish was detected on 20 December 2020, an Emergency Strategy was published in the following month to guide the first actions to multi-staff integration and practice lionfish control in the archipelago (ICMBio 2021). This was carried out by ICMBio-Noronha, with the support of ICMBio-CMEEI, to begin jointly within a month after the presentation of the MP. It is important to note that the steps followed for the approval of lionfish management project were reinforced by proper evaluation, following standardized actions for each location and which were in accordance with the national protocols (ICMBio 2019). Such procedures are useful for developing better strategies and facilitating multi-staff adhesion, resulting in the obtaining of explanatory ecological variables for IAS control.\u003c/p\u003e \u003cp\u003eFor the specimens found in Fernando de Noronha, a high physiological condition was observed and the L-W relationship, indicating high consistency between measurements. The allometric coefficient was positive, characterizing that the growth of larger individuals presents proportionally more body mass in relation to length. This scenario indicates that the lionfish population in this region exhibits growth typical of species with good energy supply and favorable body condition (Edwards et al. 2014; Ch\u0026aacute;vez-L\u0026oacute;pez et al. 2025). These factors are associated with good nutritional condition, an environment with good resource availability, and the absence of persistent energy stress (del R\u0026iacute;o et al. 2023). Substrate composition in Fernando de Noronha presented a variety of environments, in reference to the field records, with greater detection for epibenthic organisms related to corals, but also for algae and porifera, in addition to artificial structures, such as shipwrecks and anchors. Lionfish individuals are associated with a diverse set of habitats including different conditions of visibility, lightness, depth and environments (Schofield 2010). Fish behavior was noted with greater frequency for fish in resting behavior, which is consistent with the well-known low mobile behavior of lionfish (Jud and Layman 2012). In addition, we also encourage efforts to detect seasonal patterns over time. Despite the temporal series, the initial invasion of lionfish in archipelago did not allow for a relevant sampling effort to detect differences between seasons or relationships with sea conditions, which may be better indicated with a greater establishment of the species.\u003c/p\u003e \u003cp\u003eOur work corroborates with Soares et al. (2023), especially regarding collaborative network of actions for raising public awareness and developing education programs. We have provided an important baseline on the invasion of this species in new locations in South Atlantic, in addition to adequately addressing the procedures for adapting and integrating multiple teams. The local reality of Fernando de Noronha is composed of singular features, considering it is an isolated and small territory. The invasion of lionfish continues to advance to other locations in Brazil, and other difficulties may influence the control of this species, such as the frequency of diving, standardized training of multi-staff, and specific regulations. We add that each location could be delimited, based on protected areas boundary or municipalities, and communities. This makes it possible to understand local realities and apply standardized protocols through regulations. Also, new methods of control must be employed, especially in waters that are not suitable for diving, where it is possible to apply trapping techniques, such as pots and other fishing techniques (Pitt and Trott 2015). That scenario is still lacking and includes improving control methods in mesophotic and rough marine environments (Gress et al. 2017), such as the leeward sea in Fernando de Noronha.\u003c/p\u003e \u003cp\u003eIn conclusion, this research can add important perspectives for the development of national or regional regulations (e.g., ICMBio 2019; 2021; CPRH 2024), as well as the integration of international protocols, since the invasion of lionfish depends on an integrated effort (Green 2012). In addition, important advances regarding the use of captured individuals, such as gastronomic consumption, can be made regularly as a form of control. These actions should always be reinforced by educational activities, as we presented that open communication and multi-staff integration have become crucial tools for promoting an adequate strategy for lionfish control.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eConflict of interest\u003c/h2\u003e \u003cp\u003eThe authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eAuthor contributions\u003c/h2\u003e \u003cp\u003eThe project was conceptualized by RA, CBPES, LPSS, PPC, TEC and TCSG. All authors contributed to the project administration, resources, and data collection. Data analyses were performed by Lucas Penna Soares Santos. The first draft of the manuscript was written by Clara Buck Pereira do Eirado Silva and Lucas Penna Soares Santos. All authors commented on previous versions of the manuscript. All authors read and approved of the final manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgements\u003c/h2\u003e \u003cp\u003eThis work forms the coordination of research and management of ICMBio-Noronha, represented by R. Ara\u0026uacute;jo. Ethical permissions were supported by ICMBio-CMEEI (Management Project/Permission n. 10/2021 SEI n. 9919898). We thank all local agents who worked collaboratively and contributed to the lionfish management strategy in Fernando de Noronha, especially the dive instructors M. Wilson (Mike), R. Rocha (Beto), A. Gonzaga, A. Pereira (Babu), F. da Costa (Lula), J. Huss, and L. Lopes. A special strengthening was provided by Sea Paradise, represented by F. Rodrigues, the Dive Centers of \u0026Aacute;guas Claras, Atlantis Divers, Noronha Diver, Mar de Noronha, Maravista Mergulho, as well as the Image Companies of All Angle, Barracudas, Ciliares, Hidrosfera, and Natureza Viva. We appreciate all the support provided by STINAPA, especially P. Bertuol, Projeto Conserva\u0026ccedil;\u0026atilde;o Recifal (PCR), particularly P. Cipresso and L. Guilherme, PELD-ILOC, and researchers. SEMAS and CPRH established the network to control lionfish in the state of Pernambuco, with the cooperation of P. Tavares, S. Vieira, and D. Alves. To the ICMBio-Noronha Team, the Administration of the Port, Corpo de Bombeiros, visitors\u0026rsquo; guides, boat operators and community of Fernando de Noronha. We thank the reviewers who contributed to this article.\u003c/p\u003e\u003ch2\u003eData availability\u003c/h2\u003e \u003cp\u003eData will be made available on request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAirey ME, Fogg AQ, Drew JA (2023) Invasive lionfish dispersal between shallow-and deep-water habitats within coastal Floridian waters. 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In this article, we detail the management actions between 2020 and 2025 in the Fernando de Noronha archipelago, a world heritage site and biodiversity hotspot. A total of 2,660 lionfish individuals were recorded during 58 months of management, noting exponential population growth (345.9% of increase), with the number of records being higher in monitoring (n\u0026thinsp;=\u0026thinsp;2,258; focused dives) compared to routine (n\u0026thinsp;=\u0026thinsp;402; recreational dives) operations. Field detection was more related to ecological variables such as deep waters, coral substrate, weak marine currents, and resting fish behavior. High densities were generally observed at 3.9 ind./ha but increased over time and depth gradient, with a maximum value of 60 ind./ha. Individuals averaged 23.17\u0026thinsp;\u0026plusmn;\u0026thinsp;6.7 cm in total length and 238.2\u0026thinsp;\u0026plusmn;\u0026thinsp;241.6 g in weight (max.: 460 mm and 1670.0 g), presenting allometric growth (b\u0026thinsp;=\u0026thinsp;3.22) and high physiological condition (K\u0026thinsp;=\u0026thinsp;1.43). Lionfish showed high dispersion with 116 sites recorded around the archipelago and well explored areas represented 353.64 ha. Interpolation analysis reached 75,742 (CI 95% 56,202\u0026ndash;95,372) individuals regarding local population estimate. Field management proved to be effective locally in maintaining a stable temporal trend with low density averages. This work demonstrates that the use of legal and protocolary adoptions has served efficiently to stabilize and outline lionfish control through standardized actions.\u003c/p\u003e","manuscriptTitle":"Management strategy based on multi-staff integration target control of lionfish invasion on a South Atlantic oceanic island","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-02-04 18:15:33","doi":"10.21203/rs.3.rs-8626827/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2026-03-03T07:53:02+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-02-02T11:55:07+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Biological Invasions","date":"2026-01-23T23:10:59+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-01-20T08:32:09+00:00","index":"","fulltext":""},{"type":"submitted","content":"Biological Invasions","date":"2026-01-19T06:06:03+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"biological-invasions","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"binv","sideBox":"Learn more about [Biological Invasions](https://www.springer.com/journal/10530)","snPcode":"10530","submissionUrl":"https://submission.nature.com/new-submission/10530/3","title":"Biological Invasions","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"9eb8ead1-290d-4b93-84a1-9ab1b89136c3","owner":[],"postedDate":"February 4th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-02-04T18:15:33+00:00","versionOfRecord":[],"versionCreatedAt":"2026-02-04 18:15:33","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8626827","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8626827","identity":"rs-8626827","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}