Zootaxa

The genus Phakellia comprises predominantly fan-shaped deep-sea sponges, but its species diversity and boundaries remain unresolved. Here, we use an integrative approach combining morphological and molecular data to investigate the Phakellia robusta species complex and describe two new species, Phakellia pallida sp. nov. and Phakellia pusilla sp. nov. Overlap in external and spicular morphology makes these taxa challenging to identify. Molecular analyses based on mitochondrial (cox1) and nuclear (28S) markers support the recognition of the three taxa as distinct species, and reveal their close evolutionary relationship. The two new species co-occur in the NE Atlantic Ocean and the Mediterranean Sea, with overlapping depth ranges, although P. pallida sp. nov. is generally found at greater depths. Our results also suggest that several previous records of P. robusta from the Mediterranean and southern parts of the NE Atlantic may instead correspond to either P. pallida sp. nov. or P. pusilla sp. nov., indicating that its distribution requires reassessment. In addition, a group I intron previously reported in the cox1 gene of P. robusta was polymorphically detected at the same insertion site in some, but not all, individuals of P. pusilla sp. nov., but it was absent from P. pallida sp. nov. individuals. Overall, our results reveal pseudocryptic diversity within Phakellia and highlight the value of integrative approaches for resolving species boundaries and reassessing deep-sea sponge diversity and distributions.

The fieldwork was carried out between December 2023 and June 2025, with the aim, among others, of confirming the possible presence of the North African Gerbil (Gerbillus campestris) in Melilla. To this end, 56 camera trap stations were installed, each equipped with a camera trap. Of the total number of camera trap stations installed, the presence of Gerbillus campestris was detected in six of them, representing a capture success rate of 10.71%. Following the detection of the species, a specimen was captured using cage traps. For identification purposes, samples of epithelial tissue from the ear and faeces were collected. These samples were sent to the Institute of Vertebrate Biology of the Czech Academy of Sciences, Brno, Czech Republic, for analysis. There, DNA was extracted from the samples using the Innu PREP DNA Mini Kit 2.0 (IST Innuscreen GmbH, Germany) according to the manufacturer's protocol. The results obtained confirm, for the first time through records of the species, the presence of Gerbillus campestris in Melilla and, therefore, in Spain and Europe.

Ground-nesting wild bees require suitable soil conditions for their below-ground development and overwintering life stages. Understanding how soil translocation and disturbance affect individuals and the following generations of bees is important for identifying conservation measures for ground-nesting wild bees. We investigated the effects of soil translocation and disturbance on the ground-nesting solitary spring mining bee Colletes cunicularius. In a replicated controlled design, we compared the effects of soil translocation with different degrees of disturbance on the number of surviving bees and the emergence time. We found that soil translocation and disturbance reduced bee survival but did not result in complete mortality, with some individuals surviving even under severe soil disturbance. There was no evidence that soil disturbance affected the timing of spring emergence, suggesting that disturbance is unlikely to alter the timing of nest construction and, consequently, the production of the next generation. Our results indicate that translocating soil containing overwintering bees could be a promising conservation measure to compensate for habitat loss caused by land conversion.

Abstract Habitat configuration influences population connectivity and, consequently, genetic structure. River networks provide heterogeneous, hierarchically arranged environments that shift drastically from upstream to downstream. We compared congeneric species across Ephemeroptera, Plecoptera, and Trichoptera, emphasizing longitudinal replacement (upstream to downstream) and the rarely studied wet rock (hygropetric) habitats. We surveyed six rivers on the Muroto Peninsula, Japan, qualitatively sampling aquatic insects at 63 sites. Cluster analysis based on the environmental data classified surveyed sites into four clusters. We analyzed a total of 10 species from four genera that exhibit longitudinal replacement patterns within genera. Genetic analyses based on the mitochondrial cytochrome c oxidase subunit I region revealed that upstream species showed higher genetic diversity than downstream species. In contrast, species adapted to hygropetric exhibited the lowest genetic differentiation among all habitat types. A novel contribution of this study is the inclusion of hygropetric species. The surprisingly low differentiation in hygropetric species suggests high connectivity, similar to lentic species. By comparing congeneric taxa across orders within environmentally similar rivers, we reduce phylogenetic and environmental confounds, strengthening inference that habitat configuration and dispersal traits jointly shape genetic structure. These findings provide a new perspective on riverine spatial ecology and underscore the importance of microhabitat-aware comparisons for evolutionary inference.

This paper presents a comparative analysis of the floristic composition and site characteristics of four peat bogs in Bucovina, Romania: Poiana Stampei, Romanesti, Saru Dornei, and Gaina-Lucina. The research was based on phytosociological releves on 25 msq plots and direct field phenological observations, on six field visits from May to August 2026. Vegetation was characterised using the Braun Blanquet method, and floristic similarity between sites was assessed with the Sorensen and Bray Curtis indices. All four plots shared a common core of taxa characteristic of peatland vegetation: Sphagnum spp., Carex rostrata, Drosera rotundifolia, Eriophorum vaginatum, and Vaccinium species. Species richness was 13 taxa at Poiana Stampei, Romanesti, and Saru Dornei, and 12 at Gaina-Lucina. Romanesti and Saru Dornei showed the highest floristic similarity (descriptive values, not statistically tested, given a single releve per site), while Gaina-Lucina differed most markedly, not through species richness, which was similar across sites, but through species identity and through the presence of Betula nana, a glacial relict absent from the other sites. The results provide a descriptive basis for future research on the floristic composition and conservation of these habitats.

Sea turtle conservation relies largely on the identification of ecologically significant marine areas. Green sea turtles (Chelonia mydas) are resident in Aotearoa New Zealand waters, yet much of their spatial ecology remains undefined. To address this, satellite telemetry was used to map juvenile and sub-adult green sea turtle spatial movements, residency patterns, and habitat associations. Our findings provide the first evidence of individual fine-scale habitat use and long-term residency in New Zealand waters. Data will contribute to regional and global efforts to identify ecologically significant sea turtle habitat, including Important Marine Turtle Areas (IMTAs) and Blue Corridors for Turtles (BC4T), and help advance conservation efforts.

1. Protecting and restoring corals is critical to conserve marine biodiversity and ecosystem services in the Anthropocene. Knowledge of coral larval dispersal dynamics can be used to identify important sites for coral metapopulation connectivity and to spatially optimise coral conservation efforts. However, we know little regarding the larval dispersal of many coral species and our understanding of the relationships between coral larval dispersal and life-history traits (e.g., reproductive mode, colony morphology), phylogenetic relationships, and biogeography is incomplete. These knowledge gaps impede coral conservation. 2. In this research, we systematically reviewed the species, regions, and methodologies of studies of coral larval dispersal. Additionally, we conducted a meta-analysis to quantify the heterogeneity in reported coral larval duration values and identified the roles of life-history traits in driving that heterogeneity. 3. We found that various methods (experimental, biophysical modelling, and population genetics) were used to study 221 coral species across many regions. Soft and temperate corals received comparatively lower research attention. Our meta-analysis reveals that reproductive mode and colony morphology explain significant heterogeneity in larval settlement times, and that closely-related species have more similar larval settlement times. 4. Our findings suggest that reproductive mode and colony morphology could be used to predict larval duration in under-studied coral species, and that larval duration could be approximated from closely-related corals when trait data is unavailable. Our results support strategies focused on modelling differing larval durations across corals with different life-history traits to predict realised dispersal in under-studied coral species and identify conservation priority sites.

Extensive green roofs are an important component of urban green infrastructure. However, harsh rooftop conditions, including shallow substrates, high temperatures and limited water availability, considerably restrict plant selection. Although extensive green roofs are traditionally dominated by succulent species, increasing attention has been directed towards annual plants because of their rapid establishment, developmental plasticity and potential contribution to vegetation diversity. This review aimed to identify annual plant species introduced on extensive green roofs and to characterize the functional traits most frequently associated with their establishment under rooftop conditions. In addition, the review examined the geographical distribution and methodological characteristics of existing studies and identified species with potential for application under temperate climatic conditions. A total of 22 studies published between 2015 and 2025 were analyzed, comprising 100 annual species representing 29 plant families. The reviewed studies showed a clear geographical bias towards Mediterranean and arid regions, whereas temperate climates were considerably underrepresented. Among species occurring in Central Europe, ruderal life strategies, drought avoidance, efficient dispersal mechanisms, low plant height and prolonged flowering were the most frequently observed functional characteristics, while seed mass and specific leaf area (SLA) exhibited substantial interspecific variation. These findings suggest that functional traits, rather than taxonomic affiliation alone, provide a useful framework for selecting annual species for extensive green roofs. The review also highlights the need for long-term studies conducted under temperate climatic conditions to improve evidence-based plant selection and support the development of resilient and multifunctional green roof vegetation.

Fallen fleshy fruits are temporary resource patches that can support diverse but transient invertebrate assemblages, yet their successional dynamics remain poorly documented in tropical landscapes modified by human activity. We examined heterotrophic succession on fallen ripe guava fruits, Psidium guajava, in a tropical grove in Barasat, West Bengal, India. Eight fruits were placed in exposed and shaded microhabitats and followed from fresh to degraded stages over 22 days. Invertebrates were recorded using repeated scan sampling, and daily observations for each fruit were analysed for taxon richness, Shannon diversity, total peak relative abundance, community composition and indicator taxa associated with decomposition stages. Across 141 daily fruit observations, we recorded 44 invertebrate taxa. Taxon richness, Shannon diversity and total peak relative abundance were highest during fresh and pre rotten stages and declined sharply during post rotten and degraded stages. Decomposition stage was the strongest predictor of alpha diversity, resource use intensity and community composition, whereas differences between exposed and shaded microhabitats were weaker. Indicator taxa showed clear successional replacement: fresh fruits were associated with drosophilid and tephritid flies; fresh and pre rotten fruits with nitidulid beetles and ants; and later stages with collembolans, staphylinids, blattodeans and other taxa associated with detrital stages. Ants, especially Pheidole sp., were conspicuous resource users and showed guarding, hoarding, predation and pulp or seed removal behaviour, although their effects on other taxa remain observational. These results show that fallen guava fruits function as ephemeral heterotrophic resource patches, supporting rapid invertebrate turnover at fine temporal scales in a tropical grove.

The transformative impacts of global bleaching events are now a ubiquitous feature of coral reefs, often resulting in taxonomically simplified coral assemblages that can persist long-term due to ongoing ocean warming, heatwaves, and localized anthropogenic stressors. Understanding to what extent these processes shape symbiont biodiversity continues to be a major knowledge gap. Here, we examined Acropora host and symbiont communities on Kiritimati (Republic of Kiribati), seven and a half years after the 20152016 El Nino-associated marine heatwave, which caused a near total loss of the genus in the atoll's shallow reefs. We quantified Acropora cover and growth-form composition at 37 sites spanning a gradient of local human disturbance and characterised Symbiodiniaceae communities using ITS2 DNA metabarcoding from 109 coral tissue samples at 13 of the sites. Corymbose and tabular Acropora growth forms remained evenly represented across the atoll and cover declined with increasing human disturbance. Symbiodiniaceae communities were dominated by Cladocopium with 93 % of colonies associated with a single ITS2 profile. Communities were homogenous, with low diversity and insignificant differences amongst growth forms, but slight differences across disturbance levels. DNA barcoding of the PaxC intron identified the majority of host colonies as most similar to Acropora granulosa (62 %) and Acropora hyacinthus (24 %). Although dominance by a single symbiont lineage is common among Acropora species across the Pacific, the low Symbiodiniaceae diversity observed at Kiritimati contrasts with the substantial within- and among-lineage diversity documented elsewhere. The depauperate Symbiodiniaceae communities observed in this study may thus represent a lasting legacy of the 2015-2016 disturbance, with the recovering Acropora assemblage representing an early stage of community reassembly drawing from a potentially depleted environmental symbiont pool.

The identification and use of stress-tolerant coral in outplanting efforts has been proposed as a method of restoring and bolstering reef health under increasing environmental change. Prior research has focused on understanding survival in cross-reef transplantation, but evidence of long term resiliency of outplanted corals remains limited. Here, we report data at a five-year timepoint of a reciprocal transplant study of the stress-tolerant Caribbean corals Pseudodiploria strigosa and Siderastrea siderea between a nearshore and offshore site in Belize. Extreme marine heat wave conditions in August of 2024 revealed species and site-specific bleaching patterns. Through assessments of mortality, energy reserves, and chlorophyll concentrations, P. strigosa appear to be more limited in long-term acclimatization potential to the nearshore environment while S. siderea maintain acclimatization capacity displayed 17 months post transplant to nearshore. This work conveys information relevant to managers in selecting species for restoration efforts based on short-term growth and long-term resilience in native versus non-native sites, offers insight into potential mechanisms driving these trends, and takeaways on site specific management decisions for successful coral outplanting success.

Recent years have witnessed the discovery of an immense diversity of protists in almost all ecosystems on Earth. However, a major challenge consists in identifying diversity patterns and the drivers that explain the distribution of this diversity. While several works have focused on local parameters, only a few studies have addressed this question at regional scales. In this study, we characterize the diversity and distribution of Arcellinida (testate amoebae) at regional scale of a climatically contrasted country, Bulgaria. We collected 75 samples geographically evenly distributed and representative of the country's natural ecosystems and used metabarcoding to determine their diversity. Communities composition differed markedly among habitats, revealing sharp beta diversity patterns. "Mesophilic" environments such as forests and ponds were most diverse, while acidic and oligotrophic Sphagnum peatlands were poorest and hosted the highest proportion of unique taxa. Our analyses of richness-environment based on a Random Forest approach applied to all soil systems indicated that stable humidity (low silt content, low precipitation seasonality) and intermediate soil carbon stocks had a positive effect on richness. The predicted increase in continentality in the next decades in Bulgaria calls for testing climatic models to evaluate the potential risk of declining Arcellinida biodiversity.

Environmental concerns and the need to mitigate the effects of climate change have driven the adoption of renewable energy sources, such as offshore wind farms (OWFs), making it essential to assess their overlap with seabird habitats to estimate potential impacts. This study is the first to estimate the vulnerability of coastal and seabirds to OWFs at a national scale by integrating biological trait data from the literature, spatial occurrence records from citizen science platforms, and expert knowledge (elicited from 23 specialists) into established quantitative risk frameworks. Collision risk and susceptibility scores were calculated at the family and subfamily levels, as well as disturbance and displacement indices by species and susceptibility by state. The groups with the highest collision risk and susceptibility were Fregatidae (0.354; 0.486), Sulidae (0.199; 0.419), Sterninae (0.190; 0.384) and Diomedeidae (0.190; 0.416). The extreme risk for Fregatidae is driven by their specific flight characteristics, with impacts concentrated in the highly frequented southeast and northeast regions. Regarding disturbance and displacement, Limnodromus griseus exhibited the highest national score (MMA: 0.098), while Diomedea dabbenena had the highest global score (IUCN: 0.092), highlighting the critical need to integrate both regional and global conservation statuses. Spatially, the state exhibiting the highest overall vulnerability to OWF impacts was Rio Grande do Sul (23.78; 10.69), a critical hotspot characterized by high species richness and oceanographic productivity. These findings establish an indispensable national baseline for Marine Spatial Planning, reinforcing the need for targeted pre-installation assessments, the identification of sentinel species, and spatially differentiated management during environmental licensing processes.

The Peruvian Andes, and especially its slopes towards the Amazonian Basin are well-known biodiversity hotspots for plants and animals, which represent the visible part of biodiversity. However, not much is known about microeukaryotic diversity, despite the significant role protists play in the ecosystem. Based on a molecular survey of the diversity of Arcellinida (a group of amoebae well known for its environmental sensitivity and restricted geographical distributions), we estimated its diversity along an elevation gradient spanning more than 3000 meters. Surprisingly, Neotropical forest Arcellinida were less abundant and diverse than in most other extratropical biomes; nevertheless, communities differed strongly in their taxonomic composition as compared with forests in higher latitudes. Notably, sequences related to Trigonopyxis were most abundant, whereas this group occurs only marginally in extratropical settings. Elevation influenced beta diversity, showing different assembly processes occurring at different elevation belts. Canopy cover positively influenced both phylogenetic and taxonomic diversity, showing the importance of above and below ground integration in tropical forests.

Reliable morphological criteria for sexing birds without the need for invasive procedures are scientifically and ethically valuable. Smith et al (2005) reported that iris colour assigns the genetic sex of Starlings Sturnus vulgaris with 98% accuracy, and bill base colour with 100% accuracy when birds are in breeding condition, but these figures have not been tested independently. We applied their criteria to 31 captive Starlings in breeding condition, using three observers blind to genetic sex, two of them new to Starling sexing. Bill base colour correctly predicted genetic sex for all 29 birds scored, with complete observer agreement. Mean iris score was correct for 27 of 31 (87.1%), fewer than the 94 of 96 reported by Smith et al (2005) for birds in breeding condition (Fisher exact test, P = 0.031), although with so few errors in either study this comparison is imprecise. Observer agreement was high, so these errors reflect iris colouration conflicting with genetic sex rather than inconsistent scoring; bill base colour was correct for all four misassigned birds. Body mass and tarsus length classified 77% and 58% of birds correctly, their published thresholds missing in opposite directions. We conclude that bill base colour should be the primary criterion for sexing Starlings in breeding condition; the accuracy of iris colour should not be assumed to be 98%.

The endosymbiosis between tropical reef-building corals and dinoflagellate algae (family Symbiodiniaceae) underpins the healthy functioning of coral reef ecosystems. Although dominant host-symbiont pairings differ across spatial scales (e.g., bioregion, reef zone, and depth), corals often associate with multiple symbionts. Where diverse symbionts occur together, their local-scale ecology is poorly understood. Moreover, whilst coral morphological plasticity generates a structurally complex spatial habitat for endosymbiosis, the role of host geometry in facilitating symbiont co-occurrence is completely unexplored. Here, we integrated high-throughput sequencing of the ITS2 gene region with digital 3D photogrammetry to evaluate symbiont distribution within coral structural habitat and determine the influence of host morphology on symbiont community composition. Symbiont composition was characterised along a vertical gradient (coral apex to base) across 50 Hydnophora microconos colonies from Rarotonga, Cook Islands. We identified dominance by the genera Durusdinium and Cladocopium, with inter-genus co-occurrence recognised in 60% of coral colonies. The co-occurrence of two Cladocopium lineages (C3f and C3k) was validated based on congruence with the plastid minicircle psbA non-coding region (psbAncr). Geometric quantification from digital 3D reconstructions of coral colonies revealed that single symbiont taxon colonies were morphologically distinct, whilst multi-symbiont colonies exhibited a broader morphological range. Where Cladocopium-Durusdinium co-occurrence was identified, surface orientation indicated increased Durusdinium prevalence toward the base of coral colonies, corresponding to regions with the highest occlusion of modelled downwelling light. Collectively, these findings suggest that coral structural heterogeneity supports Symbiodiniaceae diversity and highlights the value of resolving microhabitat-level variation to understand symbiont co-occurrence.

Coral skeletal integrity underpins reef habitat complexity and coastal protection, yet it remains unclear whether corals persisting in chemically variable marginal habitats, such as mangrove-associated reefs, maintain sufficient mechanical resistance to withstand physical disturbance. We combined micro-computed tomography (micro-CT) morphometric analysis with compression testing to quantify skeletal microarchitecture and compressive mechanical performance of Pocillopora sp. skeletons collected across a habitat gradient at Low Isles, Great Barrier Reef, from mangrove-associated sites to exposed reef sites. Baseline skeletal properties varied among sites, with the Proximal Reef site having the densest and least porous skeletons, while mangrove-associated sites had the most porous skeletons. Similarly, Proximal Reef skeletons exhibited two-to-three-fold greater compressive stiffness and peak force than mangrove-associated skeletons. Percentage object volume was a stronger predictor of compressive stiffness than surface area-to-volume (S/V) ratio, indicating that bulk skeletal density better captured load-bearing capacity than surface architectural complexity. Structural failure mode also differed markedly among habitats: mangrove-associated skeletons underwent catastrophic fragmentation of their open, porous framework, whereas Distal and Proximal Reef skeletons failed through discrete crack-plane propagation while largely maintaining overall structural coherence. These results indicate that the physiological stress tolerance associated with mangrove habitats occurs alongside reduced mechanical resistance and more destructive failure modes, suggesting a potential trade-off between chemical stress tolerance and structural investment. Assessments of resilience in marginal coral populations should therefore consider skeletal mechanical performance alongside physiological tolerance, particularly as climate change intensifies storms and alters hydrodynamic disturbance regimes.

Understanding floral visitor dynamics along elevational gradients is essential for assessing potential implications for pollen limitation and population genetic processes in fragmented populations of cliff-dwelling alpine endemics. Physoplexis comosa (Campanulaceae) is a chasmophytic paleoendemic restricted to vertical rock faces on dolomitic and limestone bedrock from 200 to 2,800 m a.s.l. The species exhibits a specific floral morphology with fused corolla apices, basal openings for nectar access, and secondary pollen presentation, yet its functional floral visitor assemblage has remained poorly characterized across the species' elevational range. In this pilot study, we tested a two-tier camera-monitoring setup combining action cameras for diurnal monitoring with Raspberry Pi-based systems equipped with IR illumination for combined diurnal and nocturnal recording. We conducted monitoring across five elevational sites (1,480 to 2,400 m a.s.l.) in the Eastern Alps. Recorded video material was downsampled and annotated using the Addax platform alongside manual cross-validation to record physical contact events. Preliminary results confirm the technical feasibility of continuous monitoring in steep terrain and suggest possible elevational variation in diurnal visitors, alongside nocturnal activity by Geometridae moths. Camera-based observations provided higher functional resolution and revealed a different visitor assemblage than previous eDNA metabarcoding trials. This proof-of-concept establishes a methodological foundation for forthcoming multi-day monitoring campaigns and complements ongoing studies of the mating system and population-genetic analyses.

Fire-dependent or fire-influenced ecosystems harbor species with diverse response strategies, such as smoke-stimulated germination and dormancy release through heat shocks. Although these cues influence germination in many fire-prone ecosystems, their effects on seeds of woody species from tropical dry forests (TDFs) remain poorly understood. We evaluated the germination response of twelve TDF woody species to three heat-shock temperatures (80, 120, and 160C), two exposure times (3 and 5 min), and two smoke conditions (with and without smoke water), plus dry and wet controls. We also assessed the relationship between germination and functional seed traits, including permeability, seed-coat thickness, mass, and shape index. Of the eleven species that germinated, four showed broad tolerance, with no significant reduction in germination even at 160C for 5 min, which was associated with lower permeability, thicker seed coats, and greater mass. Four other species showed restricted tolerance to heat, with significant declines in germination at higher temperatures or longer exposure times, and were associated with higher permeability, thinner seed coats, and lower seed mass. The remaining three species showed heat-stimulated germination between 80 and 120C and tended to have lower shape-index values. Smoke treatments significantly increased germination in three species, Ceiba pentandra, Guazuma ulmifolia, and Samanea saman. In the remaining species, significant differences were either absent or attributable to moisture or imbibition effects rather than to smoke itself. Overall, germination responses to heat were associated with variation in functional seed traits, whereas responses to smoke were less widespread and showed a less consistent functional pattern.

Reef fish assemblages in the Veracruz Reef System National Park (VRSNP) are strongly structured along a pronounced north-south gradient, where environmental conditions, urban proximity, and protection levels interact to shape functional diversity and ecosystem functioning. This gradient creates contrasting ecological conditions between northern and southern reefs, providing a natural framework for evaluating how spatial variation influences resilience. Using a stationary visual census across ten reefs distributed latitudinally, we quantified functional entities based on six ecological traits and calculated functional richness, redundancy, over-redundancy, vulnerability, specialization, and originality. A two-way permutational ANOVA and redundancy analysis were used to detect spatial and temporal patterns and to link functional indices with environmental features. The southern sector displayed the most intense functional shifts. Nearshore southern reefs, strongly exposed to sedimentation, nutrient loading, and physical disturbance, exhibited reduced functional richness, fewer functional entities, and high functional vulnerability, indicating strong environmental filtering. Over time, these reefs shifted toward generalist-dominated assemblages with rising redundancy, suggesting functional convergence. In contrast, southern offshore reefs supported higher taxonomic and functional richness, buffered by greater habitat complexity and distance from shore, although they showed declining redundancy, likely reflecting sensitivity to regional climatic disturbances. In the northern sector, the gradient was moderated by effective protection, which reshaped functional configurations. Nearshore protected reefs such as Isla Sacrificios and Isla de Enmedio maintained high redundancy, low vulnerability, and high biomass of large-bodied fishes, resembling offshore systems rather than disturbed southern nearshore reefs. The northern offshore reef (Isla Verde) shared some of these attributes but exhibited temporal declines in richness, signaling emerging vulnerabilities along the gradient. Functional originality and specialization aligned with benthic states across the gradient, particularly where macroalgal or calcareous substrates prevailed. Overall, the north-south gradient is the primary axis of functional organization in VRSNP, revealing that resilience emerges from the interplay among reef geomorphology, urban influence, and governance effectiveness.