Evolutionary Ecology Research Articles

Ecology, evolution, and conservation of Ethiopia’s biodiversity

Ethiopia is home to one of the richest and most unique assemblages of fauna and flora on the African continent. Contained within its borders are two major centers of endemism, the mesic Roof of Africa (also known as the Ethiopian Highlands) and the arid Horn of Africa, resulting from the country’s varied topography and consequent geographic isolation. These centers of endemism are crucial to global conservation as evidenced by their classification within the Eastern Afromontane and Horn of Africa biodiversity hotspots, respectively. Ethiopia’s diverse ecosystems and the biodiversity they contain are increasingly threatened by climate change and the growing impacts of Africa’s second largest human and largest livestock populations. In this paper, we focus on several key areas of recent and ongoing research on Ethiopian biodiversity that have broadened our understanding of nature and its conservation in Africa. Topics explored include the behavioral ecology of Ethiopia’s large social mammals, the ecology and conservation of its unique coffee forests, and Ethiopian approaches to community conservation, fortress conservation, and nature-based solutions. We also highlight the increasing prominence of Ethiopian scientists in studies of the country’s biodiversity in recent decades. We suggest promising avenues for future research in evolutionary biology, ecology, systematics, and conservation in Ethiopia and discuss how recent and ongoing work in Ethiopia is helping us better understand and conserve nature in the human-dominated landscapes of Africa and other tropical regions today.

Lessons from comparative primatology for understanding trait covariation and diversity in evolutionary ecology

ABSTRACT Given the position of humans in the tree of life, comparative research on non‐human primates has attracted the interest of researchers in biology, medical sciences, anthropology, psychology, and sociology. Covariation of species' phenotypes has been of particular interest. Learning from the historical development of comparative research with primates should thus be particularly valuable for evolutionary ecology and to improve understanding of phenotypic integration and diversity. Such learning would also help identify knowledge gaps, disputed questions, and new avenues of both basic and applied research in relation to the evolution of primate features and the conservation of our close relatives. We conducted a historical assessment through a non‐systematic review and a systematic review, focusing on how the integration of different research lines in evolutionary ecology focused on primate phenotypic covariation unfolded throughout the 20th Century. The non‐systematic review allowed us to reconstruct the history of the discipline from its earliest origins, when bibliometric assessments were more limited in scope, and to identify the most appropriate keywords for the systematic review. We employed a standard protocol for the systematic review, applying two complementary analyses: co‐occurrence of keywords and bibliographic coupling of references. These analyses described the development of the conceptual and intellectual structures of comparative primatology from 1966 to 2020. By identifying the most influential researchers and concept interrelations, we highlight primate phenotypes critical for the development of the discipline (in particular, brain and body sizes and behavioural patterns), showcasing the reach of these investigations for evolutionary ecology. Overall, our findings emphasise the crucial role that comparative primatology has played in developing the study of phenotypic integration and the very onset of phylogenetic comparative methods.

Editorial: Nesting in reptiles: Natural and anthropogenic threats and evolutionary responses

EDITORIAL article Front. Ecol. Evol., 06 December 2022Sec. Behavioral and Evolutionary Ecology Volume 10 - 2022 | https://doi.org/10.3389/fevo.2022.1103193

Ecology, Not Host Phylogeny, Shapes the Oral Microbiome in Closely Related Species.

Host-associated microbiomes are essential for a multitude of biological processes. Placed at the contact zone between external and internal environments, the little-studied oral microbiome has important roles in host physiology and health. Here, we investigate the roles of host evolutionary relationships and ecology in shaping the oral microbiome in three closely related gorilla subspecies (mountain, Grauer's, and western lowland gorillas) using shotgun metagenomics of 46 museum-preserved dental calculus samples. We find that the oral microbiomes of mountain gorillas are functionally and taxonomically distinct from the other two subspecies, despite close evolutionary relationships and geographic proximity with Grauer's gorillas. Grauer's gorillas show intermediate bacterial taxonomic and functional, and dietary profiles. Altitudinal differences in gorilla subspecies ranges appear to explain these patterns, suggesting a close connection between dental calculus microbiomes and the environment, likely mediated through diet. This is further supported by the presence of gorilla subspecies-specific phyllosphere/rhizosphere taxa in the oral microbiome. Mountain gorillas show a high abundance of nitrate-reducing oral taxa, which may promote adaptation to a high-altitude lifestyle by modulating blood pressure. Our results suggest that ecology, rather than evolutionary relationships and geographic distribution, shape the oral microbiome in these closely related species.

Pollinator foraging tactics have divergent consequences for the mating system of a tropical plant.

Resolving the consequences of pollinator foraging behaviour for plant mating systems is a fundamental challenge in evolutionary ecology. Pollinators may adopt particular foraging tactics: complete trapline foraging (repeated movements along a fixed route), sample-and-shift trapline foraging (a variable route that incorporates information from previous experiences) and territorial foraging (stochastic movements within a restricted area). Studies that integrate these pollinator foraging tactics with plant mating systems are generally lacking. We investigate the consequences of particular pollinator foraging tactics for Heliconia tortuosa. We combine parentage and sibship inference analysis with simulation modelling to: estimate mating system parameters; infer the foraging tactic adopted by the pollinators; and quantify the impact of pollinator foraging tactics on mating system parameters. We found high outcrossing rates, ubiquitous multiple paternity and a pronounced departure from near-neighbour mating. We also found that plants repeatedly receive pollen from a series of particular donors. We infer that the pollinators primarily adopt complete trapline foraging and occasionally engage in sample-and-shift trapline foraging. This enhances multiple paternity without a substantial increase in near-neighbour mating. The particular pollinator foraging tactics have divergent consequences for multiple paternity and near-neighbour mating. Thus, pollinator foraging behaviour is an important driver of the ecology and evolution of plant mating systems.

Hosts, microbiomes, and the evolution of critical windows.

The absence of microbial exposure early in life leaves individuals vulnerable to immune overreaction later in life, manifesting as immunopathology, autoimmunity, or allergies. A key factor is thought to be a "critical window" during which the host's immune system can "learn" tolerance, and beyond which learning is no longer possible. Animal models indicate that many mechanisms have evolved to enable critical windows, and that their time limits are distinct and consistent. Such a variety of mechanisms, and precision in their manifestation suggest the outcome of strong evolutionary selection. To strengthen our understanding of critical windows, we explore their underlying evolutionary ecology using models encompassing demographic and epidemiological transitions, identifying the length of the critical window that would maximize fitness in different environments. We characterize how direct effects of microbes on host mortality, but also indirect effects via microbial ecology, will drive the optimal length of the critical window. We find that indirect effects such as magnitude of transmission, duration of infection, rates of reinfection, vertical transmission, host demography, and seasonality in transmission all have the effect of redistributing the timing and/or likelihood of encounters with microbial taxa across age, and thus increasing or decreasing the optimal length of the critical window. Declining microbial population abundance and diversity are predicted to result in increases in immune dysfunction later in life. We also make predictions for the length of the critical window across different taxa and environments. Overall, our modeling efforts demonstrate how critical windows will be impacted over evolution as a function of both host-microbiome/pathogen interactions and dispersal, raising central questions about potential mismatches between these evolved systems and the current loss of microbial diversity and/or increases in infectious disease.

Ediacaran Corumbella has a cataphract calcareous skeleton with controlled biomineralization.

Corumbella is a terminal Ediacaran tubular, benthic fossil of debated morphology, composition, and biological affinity. Here, we show that Corumbella had a biomineralized skeleton, with a bilayered construction of imbricated calcareous plates and rings (sclerites) yielding a cataphract organization, that enhanced flexibility. Each sclerite likely possessed a laminar microfabric with consistent crystallographic orientation, within an organic matrix. Original aragonitic mineralogy is supported by relict aragonite and elevated Sr (mean= ca. 11,800 ppm in central parts of sclerites). In sum, the presence of a polarisation axis, sclerites with a laminar microfabric, and a cataphract skeletal organization reminiscent of early Cambrian taxa, are all consistent with, but not necessarily indicative of, a bilaterian affinity. A cataphract skeleton with an inferred complex microstructure confirms the presence of controlled biomineralization in metazoans by the terminal Ediacaran, and offers insights into the evolution of development and ecology at the root of the 'Cambrian radiation'.

Species delineation and genetic structure of two Chaerephon species (C. pusillus and C. leucogaster) on Madagascar and the Comoro archipelago.

Cryptic species diversity is known to be common in bats but remains challenging to study in these mammals, whose natural history traits render their sampling and monitoring challenging. For these animals, indirect genetic approaches provide a powerful tool to gain insight into the evolutionary history and ecology of cryptic bat species. The speciation history of the polyphyletic Chaerephon pumilus species group (Molossidae) is poorly understood, including those found on western Indian Ocean islands. Two species in this complex have been identified in the Comoros: C. pusillus and C. leucogaster. Here, we aim to genetically characterize these two species and investigate their spatial population genetic structure. Analyzing five nuclear microsatellite markers from 200 individuals and one mitochondrial DNA gene (Cyt-b) from 161 (out of the 200) individuals sampled on Madagascar and the Comoros, our findings indicated that these species are genetically differentiated. We observed mitonuclear discordance in numerous individuals (33% of the 161 mtDNA-sequenced individuals). Based on ABC analyses, we found that this pattern could potentially be the result of asymmetric introgressive hybridization from C. leucogaster to C. pusillus and calls for further studies on the demographic history of these species. Moreover, at the intra-specific level, analyses of the microsatellite loci suggested the evidence of a more pronounced, although weak, geographically based genetic structure in C. pusillus than in C. leucogaster. Altogether, our findings provide preliminary insights into the eco-evolutionary aspects of this species complex and warrant further research to understand hybridization dynamics and mechanisms responsible for mitonuclear discordance.

Spatial scales of competition and a growth-motility trade-off interact to determine bacterial coexistence.

The coexistence of competing species is a long-lasting puzzle in evolutionary ecology research. Despite abundant experimental evidence showing that the opportunity for coexistence decreases as niche overlap increases between species, bacterial species and strains competing for the same resources are commonly found across diverse spatially heterogeneous habitats. We thus hypothesized that the spatial scale of competition may play a key role in determining bacterial coexistence, and interact with other mechanisms that promote coexistence, including a growth-motility trade-off. To test this hypothesis, we let two Pseudomonas putida strains compete at local and regional scales by inoculating them either in a mixed droplet or in separate droplets in the same Petri dish, respectively. We also created conditions that allow the bacterial strains to disperse across abiotic or fungal hyphae networks. We found that competition at the local scale led to competitive exclusion while regional competition promoted coexistence. When competing in the presence of dispersal networks, the growth-motility trade-off promoted coexistence only when the strains were inoculated in separate droplets. Our results provide a mechanism by which existing laboratory data suggesting competitive exclusion at a local scale is reconciled with the widespread coexistence of competing bacterial strains in complex natural environments with dispersal.

Towards a Model of Urban Evolution—Part III: Rules of Evolution

This paper develops a formal model of urban evolution in terms of (1) sources of variations; (2) principles of selection; and (3) mechanisms of retention. More specifically, regarding (1) it defines local and environmental sources of variation and identifies some of their generative processes, such as recombination, migration, mutation, extinction, and transcription errors. Regarding (2), it outlines a series of selection processes as part of an evolutionary ecology of urban forms, including density dependence, scope dependence, distance dependence, content dependence, and frequency dependence. Regarding (3), it characterizes retention as a combination of absorption and restriction of novel variants, defines mechanisms by which these can occur, including longevity, fidelity, and fecundity, and specifies how these processes issue in trajectories define by properties such as stability, pace, convergence, and divergence. A conclusion reviews the effort and looks forward to computer simulation and data-driven applications, as well as focused theoretical extensions of parts of the model. This paper builds on Parts I and II and is part of the Urban Genome Project, about which more information can be found here.

Study on the direction effect of terrane accretive orogeny and biological evolution during the Jinning–Chengjiang period in the northwestern margin of the Yangtze block

In the various stages from life chemistry to biological evolution, major geological events are usually accompanied by the extinction of species and the gestation of new life, which are important factors that determine the direction of ecological evolution. The study of the terrane accretive orogeny during the Jinning–Chengjiang period has had a great impact on the crustal rock composition, the atmospheric environment and the water circulation system, etc. The living environment of the primitive biological community has undergone tremendous changes; for example, the crust produced a superimposed contraction and extensional rift effect, resulting in collisional magmatic activity and strong earthquakes, and a large number of species became extinct. In turn, biological activity and organic matter also contributed significantly to migration. Given the periodic cycle of geological processes, and being fully utilized by nature, the origin and evolution of life grow in a spiral. Based on the analysis of the petrochemical composition characteristics of metamorphic volcanic–sedimentary rock series and intermediate-acid intrusive rocks in Pengguan terrane, this paper proposes that the subduction mechanism of oceanic crust in the study area is formed by the NW subduction of the Longmenshan–Anning oceanic crust on the SE side. The relationship between the average change of chemical composition of the diabase in the Masongling and Xinkaidong areas and around the world, ion radius and compressibility, shows that when the element ion compressibility is less than 4.46 compared with the average world value, the oxide content of Si, Ti, Al and P is markedly lower, while when it is equal to or greater than 4.46, the oxide content of Fe, Mn, Mg and K increases, as opposed to that of Ca and Na.

Importance of interindividual interactions in eco-evolutionary population dynamics: The rise of demo-genetic agent-based models.

The study of eco-evolutionary dynamics, that is of the intertwinning between ecological and evolutionary processes when they occur at comparable time scales, is of growing interest in the current context of global change. However, many eco-evolutionary studies overlook the role of interindividual interactions, which are hard to predict and yet central to selective values. Here, we aimed at putting forward models that simulate interindividual interactions in an eco-evolutionary framework: the demo-genetic agent-based models (DG-ABMs). Being demo-genetic, DG-ABMs consider the feedback loop between ecological and evolutionary processes. Being agent-based, DG-ABMs follow populations of interacting individuals with sets of traits that vary among the individuals. We argue that the ability of DG-ABMs to take into account the genetic heterogeneity-that affects individual decisions/traits related to local and instantaneous conditions-differentiates them from analytical models, another type of model largely used by evolutionary biologists to investigate eco-evolutionary feedback loops. Based on the review of studies employing DG-ABMs and explicitly or implicitly accounting for competitive, cooperative or reproductive interactions, we illustrate that DG-ABMs are particularly relevant for the exploration of fundamental, yet pressing, questions in evolutionary ecology across various levels of organization. By jointly modelling the effects of management practices and other eco-evolutionary processes on interindividual interactions and population dynamics, DG-ABMs are also effective prospective and decision support tools to evaluate the short- and long-term evolutionary costs and benefits of management strategies and to assess potential trade-offs. Finally, we provide a list of the recent practical advances of the ABM community that should facilitate the development of DG-ABMs.

Transition model for the hermaphroditism-dioecy continuum in higher plants

Dioecy, characterized by the presence of distinct male and female individuals in the breeding population, has evolved independently numerous times, suggesting that a simple common mechanism may prevail behind the evolution of dioecy from hermaphroditism. In the present study, a floral sex allocation model is demonstrated for the presence of a hermaphroditism-dioecy continuum in higher plants using an N-player game model. In the continuum, we pay attention to the emergence of gynodioecy in which both hermaphroditic and female individuals coexist in the breeding population, and trioecy which includes hermaphroditic, male and female individuals in the breeding population, as these have often been considered as an intermediate step in the evolution of dioecy from hermaphroditism and vice versa. In the N-player game model, we assumed a local breeding population containing N players with different reproductive investment, and there is a trade-off between the reproductive investment to male function and female function at individual level. Furthermore, we focused on the uncertainty of pollen transportation, as dioecy is often linked to both the unreliable pollen vectors and harsh environment for pollinators. The pollen transport efficiency was explained by the probability of the pollen hitting ovules or stigmata. In the model, setting hypothetical populations with different population sizes, we showed the Nash solutions for dioecy appeared at a smaller pollen transport efficiency than for hermaphroditism in all hypothetical populations. Since the low pollen transport efficiency equals the low reliability of pollen vectors, the result is comparable with the pollinators of real dioecy such as water, wind and opportunistic species. The Nash solution for gynodioecy and trioecy was obtained at the transition zone between dioecy and hermaphroditism in the continuum. The number of male and female individuals in dioecy were almost stable in the range of low pollen transport efficiency. If the number of males decreased with increasing pollen transport efficiency, hermaphroditic individuals emerged in the population (establishment of trioecy). Gynodioecy established as soon as no males were left in the population. Finally, hermaphroditism established at the time no females were left in the population. These results imply that pollen transport efficiency played an important role of obtaining a variety of Nash solutions in the model. Though the Darwinian paradigm in evolutionary ecology of dioecy is based on the various aspects of the interaction between environment and species-specific characters, our model is composed of numerical interactions between and within individuals. In this paper, we argue that the association between the continuum and the pollen transport efficiency is reflected in the real world.

The opportunity for selection: A slippery concept in ecology and evolution.

Natural selection can only occur if individuals differ in fitness. For this reason, the variance in relative fitness has been equated with the 'opportunity for selection' ( ), which has a long, albeit somewhat controversial, history. In this paper we discuss the use/misuse of and related metrics in evolutionary ecology. The opportunity is only realised if some fraction of is caused by trait variation. Thus, does not imply that selection is occurring, as sometimes erroneously assumed, because all fitness variation could be independent of phenotype. The selection intensity on any given trait cannot exceed , but this upper limit will never be reached because (a) stochastic factors always affect fitness, and (b) there might be multiple traits under selection. The expected magnitude of the stochastic component of is negatively correlated with mean fitness. Uncertainty in realised is also larger when mean fitness or population/sample size are low. Variation in across time or space thus can be dominated (or solely driven) by variation in the strength of demographic stochasticity. We illustrate these points using simulations and empirical data from a population study on great tits Parus major. Our analysis shows that the scope for fecundity selection in the great tits is substantially higher when using annual number of recruits as the fitness measure, rather than fledglings or eggs, even after adjusting for the dependence of on mean fitness. This suggests nonrandom survival of juveniles across families between life stages. Indeed, previous work on this population has shown that offspring recruitment is often nonrandom with respect to clutch size and laying date of parents, for example. We conclude that one cannot make direct inferences about selection based on fitness data alone. However, examining variation in (the opportunity for fecundity selection adjusted for mean fitness) across life stages, years or environments can offer clues as to when/where fecundity selection might be strongest, which can be useful for research planning and experimental design.

Mark-recapture of microorganisms.

Fundamental questions in ecology and evolution require the quantitative tracking of individuals across space and time. For more than a century, mark-recapture techniques have been used to estimate demographic events and population dynamics of animal and plant species. Recent developments in molecular barcoding allow for the monitoring of microorganisms while meeting many of the key assumptions of traditional mark-recapture studies. Beyond applications related to the biosecurity of human pathogens, molecular barcoding is emerging as a cost-effective, versatile, and scalable tool to quantify species interactions, biogeographic ranges, and evolutionary dynamics in simple and complex communities. Although there are logistical and ethical issues related to the release of barcoded organisms, mark-recapture provides opportunities to test theory and better understand the evolutionary ecology of microorganisms in the wild.

Intraspecific diversity of threespine stickleback (Gasterosteus aculeatus) populations in eastern Canada

The threespine stickleback (Gasterosteus aculeatus) is a small, mesopredatory fish that is widespread in coastal regions of the northern hemisphere. Although studied extensively as a model organism in evolutionary biology, behavioral ecology, genomics, and numerous related subfields, this research relies heavily on populations from the Pacific coastal regions of North America and Asia, and those of northern Europe. However, based on the morphology of some western Atlantic populations, the different ecological context, and the evolutionary history of the species, not all of the knowledge gained from Pacific and European lineages is likely to be fully transferrable to the populations of North America’s East Coast. Nevertheless, work in eastern Canada does suggest high levels of intraspecific phenotypic diversity and local adaptation, though much of this diversity may be under threat from climate change, altered land use patterns, and introduced species. These factors warrant a research program focused on broad sampling of previously identified populations, identifying previously undocumented populations, determining whether there are unique genetic mechanisms underlying the unusual trait combinations present in the region, and exploring novel community interactions. Such a research program would facilitate future studies in ecology, conservation, and evolution by documenting phenotypic changes and establish baselines for future work. Because the work on nearshore marine threespine stickleback populations is sparse in the western Atlantic, I focus here on freshwater populations—with the exception of a brief discussion of the “white” stickleback populations of Nova Scotia—but this is not to suggest that nearshore marine populations are not phenotypically diverse.

The mutual history of Schlegel's Japanese gecko (Reptilia: Squamata: Gekkonidae) and humans inscribed in genes and ancient literature.

Knowing how the present distribution of organisms was formed is an essential issue in evolutionary ecology. Recently, the distribution of organisms on Earth has been significantly changed by human-mediated dispersal due to globalization. Therefore, significant attention has been paid to such processes. However, although humankind has taken considerable time to achieve modernization, the impact of ancient human activity on ecosystems has not yet been thoroughly studied. We hypothesized that ancient urban development and transitions had a non-negligible effect on species distribution. Inferring the impact of past human activity on ecosystems from ancient literature and verifying that impact by genetic analysis and human history is an effective means of tackling this problem. As geckos, a popular neighbor of human dwellings, are good material for this model, we performed this combination approach using Schlegel's Japanese gecko, Gekko japonicus. We show that G. japonicus migrated from China to the western Japanese archipelago before Christ. The gecko species dispersed itself from western to eastern the archipelago on a time scale of thousands of years. There are many synchronizations between the dispersal history of G. japonicus and the historical development of human society. It is suggested by such synchronizations that humans have influenced the distribution of G. japonicus many times throughout its dispersal history.

Moving past the challenges and misconceptions in urban adaptation research.

Although the field of urban evolutionary ecology has recently expanded, much progress has been made in identifying adaptations that arise as a result of selective pressures within these unique environments. However, as studies within urban environments have rapidly increased, researchers have recognized that there are challenges and opportunities in characterizing urban adaptation. Some of these challenges are a consequence of increased direct and indirect human influence, which compounds long-recognized issues with research on adaptive evolution more generally. In this perspective, we discuss several common research challenges to urban adaptation related to (1) methodological approaches, (2) trait-environment relationships and the natural history of organisms, (3) agents and targets of selection, and (4) habitat heterogeneity. Ignoring these challenges may lead to misconceptions and further impede our ability to draw conclusions regarding evolutionary and ecological processes in urban environments. Our goal is to first shed light on the conceptual challenges of conducting urban adaptation research to help avoid the propagation of these misconceptions. We further summarize potential strategies to move forward productively to construct a more comprehensive picture of urban adaptation, and discuss how urban environments also offer unique opportunities and applications for adaptation research.

How public can public goods be? Environmental context shapes the evolutionary ecology of partially private goods.

The production of costly public goods (as distinct from metabolic byproducts) has largely been understood through the realization that spatial structure can minimize losses to non-producing "cheaters" by allowing for the positive assortment of producers. In well-mixed systems, where positive assortment is not possible, the stable production of public goods has been proposed to depend on lineages that become indispensable as the sole producers of those goods while their neighbors lose production capacity through genome streamlining (the Black Queen Hypothesis). Here, we develop consumer-resource models motivated by nitrogen-fixing, siderophore-producing bacteria that consider the role of colimitation in shaping eco-evolutionary dynamics. Our models demonstrate that in well-mixed environments, single "public goods" can only be ecologically and evolutionarily stable if they are partially privatized (i.e., if producers reserve a portion of the product pool for private use). Colimitation introduces the possibility of subsidy: strains producing a fully public good can exclude non-producing strains so long as the producing strain derives sufficient benefit from the production of a second partially private good. We derive a lower bound for the degree of privatization necessary for production to be advantageous, which depends on external resource concentrations. Highly privatized, low-investment goods, in environments where the good is limiting, are especially likely to be stably produced. Coexistence emerges more rarely in our mechanistic model of the external environment than in past phenomenological approaches. Broadly, we show that the viability of production depends critically on the environmental context (i.e., external resource concentrations), with production of shared resources favored in environments where a partially-privatized resource is scarce.

Stranding events of drift organisms (Arribadas) in southern Brazil and the spread of invasive bryozoan in South America.

The accumulation of drift organisms (Arribadas, constituted by benthic microalgae, macroalgae and bryozoans) on Central Beach in Balneário Camboriú (SC) has drawn attention since the early 2000s. However, historical surveys suggest an ancient phenomenon that has been increasing in intensity, promoted by urbanization and the eutrophication of the bay in recent years, leading to changes in the taxonomic composition. Previously, these Arribadas were comprised of two species of benthic microalgae (Amphitetras antediluviana and Biddulphia biddulphiana) and the bryozoan Arbocuspis ramosa. However, since 2019, a substantial increase in biomass has been observed on the beach and the dominance of the macroalgae Bryopsis plumosa strongly suggests an increase in the load of organic matter and nutrients in the bay. Recently (2022) the presence of a new invasive bryozoan species (Amathia alternata) was detected, highlighting the need to continue investigating the Arribadas to monitor the ecological evolution of this process.