Darwin's feathers: Eco-evolutionary biology, predictions and policy.

The scientific community is often asked to predict the future state of the environment and, to do so, the structure (biodiversity) and the functions (ecosystem functioning) of the investigated systems must be described and understood. In his "handful of feathers" metaphor, Charles Darwin explained the difference between simple and predictable systems, obeying definite laws, and complex (and unpredictable) systems, featured by innumerable components and interactions among them. In order not to waste efforts in impossible enterprises, it is crucial to ascertain if accurate predictions are possible in a given domain, and to what extent they might be reliable. Since ecology and evolution (together forming "natural history") deal with complex historical systems that are extremely sensitive to initial conditions and to contingencies or 'black swans', it is inherently impossible to accurately predict their future states. Notwithstanding this impossibility, policy makers are asking the community of ecological and evolutionary biologists to predict the future. The struggle for funding induces many supposed naturalists to do so, also because other types of scientists (from engineers to modellers) are keen to sell predictions (usually in form of solutions) to policy makers that are willing to pay for them. This paper is a plea for bio-ecological realism. The "mission" of ecologists and evolutionary biologists (natural historians) is not to predict the future state of inherently unpredictable systems, but to convince policy makers that we must live with uncertainties. Natural history, however, can provide knowledge-based wisdom to face the uncertainties about the future. Natural historians produce scenarios that are of great help in figuring out how to manage our relationship with the rest of nature.

ß-diversity reveals ecological connectivity patterns underlying marine community recovery: Implications for conservation.

As ß-diversity can be seen as a proxy of ecological connections among species assemblages, modeling the decay of similarity in species composition at increasing distance may help elucidate spatial patterns of connectivity and local- to large-scale processes driving community assembly within a marine region. This, in turn, may provide invaluable information for setting ecologically coherent networks of marine protected areas (MPAs) in which protected communities are potentially interrelated and can mutually sustain against environmental perturbations. However, field studies investigating changes in ß-diversity patterns at a range of spatial scales and in relation to disturbance are scant, limiting our understanding of how spatial ecological connections among marine communities may affect their recovery dynamics. We carried out a manipulative experiment simulating a strong physical disturbance on subtidal rocky reefs at several locations spanning >1000 km of coast in the Adriatic Sea (Mediterranean Sea) and compared ß-diversity patterns and decay of similarity with distance and time by current transport between undisturbed and experimentally disturbed macrobenthic assemblages to shed light on connectivity processes and scales involved in recovery. In contrast to the expectation that very local-scale processes, such as vegetative regrowth and larval supply from neighboring undisturbed assemblages, might be the major determinants of recovery in disturbed patches, we found that connectivity mediated by currents at larger spatial scales strongly contributed to shape community reassembly after disturbance. Across our study sites in the Adriatic Sea, ß-diversity patterns suggested that additional protected sites that matched hotspots of propagule exchange could increase the complementarity and strengthen the ecological connectivity throughout the MPA network. More generally, conditional to habitat distribution and selection of sites of high conservation priority (e.g., biodiversity hotspots), setting network internode distance within 100-150 km, along with sizing no-take zones to cover at least 5 km of coast, would help enhance the potential connectivity of Mediterranean subtidal rocky reef assemblages from local to large scale. These results can help improve conservation planning to achieve the goals of promoting ecological connectivity within MPA networks and enhancing their effectiveness in protecting marine communities against rapidly increasing natural and anthropogenic disturbances.

Setting thresholds is not enough: Beach litter as indicator of poor environmental status in the southern Adriatic Sea.

This study deals with the issue of beach litter pollution in the context of the Descriptor 10 of the Marine Strategy Framework Directive Good Environmental Status of EU waters and Ecological objective 10, Common indicator 22 of IMAP. Analyses of the amount, distribution and categorization of beach litter were conducted on nine beaches during 108 surveys covering the area of 206.620 m2 in Albania, Italy and Montenegro. Our findings showed that the level of beach litter pollution on south Adriatic beaches is significantly above the adopted threshold values, with a median item numbers of 327, 258 and 234 per 100 m of beach stretch for Albania, Italy and Montenegro, respectively. It can be concluded that, when it comes to beach litter pollution, GES has not been achieved. Given the defined baseline and threshold values at the EU level, the process of reducing the total amount of marine litter in southern Adriatic Sea will be very challenging and needs urgent and specific actions.

Italian natural history museums need specimen digitization and much more: a reply to Benvenuti et al.

We reply to the comments made by Benvenuti et al. (2022) about our paper on the Italian natural history museums and scientific collections and the need of a centralized hub and repository. While agreeing that digitization is a useful tool to valorize each museum and collection, we still believe that the suggestion of a centralized hub is valid and necessary. This would largely help in boosting coordination among museums, sharing personnel and resources, and in providing a place to deposit scientific collections that do not fit the scope of smaller museums.

Reconnecting research and natural history museums in Italy and the need of a national collection biorepository.

In Italy, differently from other countries, a national museum of natural history is not present. This absence is due, among other reasons, to its historical political fragmentation up to 1870, which led to the establishment of medium-sized museums, mostly managed by local administrations or universities. Moreover, a change of paradigm in biological research, at the beginning of the 20th century, contributed to privilege experimental studies in universities and facilitated the dismissal of descriptive and exploratory biology, which formed the basis of the taxonomic research carried out by natural history museums. Consequently, only a few museums have a provision of curatorial staff, space and material resources adequate to maintain their original mission of discovering the natural world, by conducting a regular research activity accompanied by field campaigns. The creation of a national research centre for the study of biodiversity, facilitating interconnections among the existing natural history museums could be a solution and is here supported, together with a centralised biorepository to host collections and vouchers, to the benefit of current and future taxonomic research and environmental conservation. Such an institution should find place and realisation within the recently proposed National Biodiversity Future Center (NBFC) planned within the National Plan of Recovery and Resilience (PNRR). Pending upon the creation of this new national centre, a network among the existing museums should coordinate their activities.

The future ocean we want.

Backcasting involves the design of a desirable future that is not simply predicted with forecasts being, instead, proactively aimed at with effective action. So far, all initiatives towards sustainability failed, probably due to lack of investments in the acquisition of knowledge on the structure and the function of natural systems (i.e. biodiversity and ecosystem functioning), and to the reliance on models and estimates based on incomplete data.

Mediterranean rocky reefs in the Anthropocene: Present status and future concerns.

Global change is striking harder and faster in the Mediterranean Sea than elsewhere, where high levels of human pressure and proneness to climate change interact in modifying the structure and disrupting regulative mechanisms of marine ecosystems. Rocky reefs are particularly exposed to such environmental changes with ongoing trends of degradation being impressive. Due to the variety of habitat types and associated marine biodiversity, rocky reefs are critical for the functioning of marine ecosystems, and their decline could profoundly affect the provision of essential goods and services which human populations in coastal areas rely upon. Here, we provide an up-to-date overview of the status of rocky reefs, trends in human-driven changes undermining their integrity, and current and upcoming management and conservation strategies, attempting a projection on what could be the future of this essential component of Mediterranean marine ecosystems.

Mission possible: Holistic approaches can heal marine wounds.

Ocean sciences comprise a vast array of disciplines ranging from physics to socio-economics. The various approaches compete with each other for visibility, rather than cooperate and join forces. Communication beyond the science journals tends to focus on charismatic species and habitats (the ohhh tactics, aimed at provoking wonder) that does not result in the full perception (the ahhh strategy) of the role of ocean sciences for our well-being. Furthermore, natural sciences fail to establish the logical primacy of natural laws over social and economic laws, even though society and the economy cannot exist without the rest of the environment. Sustainability is universally recognized as a stringent priority, but it gives prevalence to economic and social values, relegating natural phenomena to a secondary role: natural assets are evaluated with measures of economics (in monetary terms) giving primacy to economic laws over ecological laws. The solution to these problems resides in the cultural evolution of that part of the scientific community that recognizes the necessity of evolving from reductionistic tactics to a holistic strategy, with a timely integration of ocean sciences and technologies and of their collaboration with social and economic sciences.

The Cells of Ecosystem Functioning: Towards a holistic vision of marine space.

Marine space is three dimensional, the turnover of life forms is rapid, defining a fourth dimension: time. The definition of ecologically significant spatial units calls for the spatio-temporal framing of significant ecological connections in terms of extra-specific (biogeochemical cycles), intra-specific (life cycles), and inter-specific (food webs) fluxes. The oceanic volume can be split in sub-systems that can be further divided into smaller sub-units where ecosystem processes are highly integrated. The volumes where oceanographic and ecological processes take place are splittable into hot spots of ecosystem functioning, e.g., upwelling currents triggering plankton blooms, whose products are then distributed by horizontal currents, so defining Cells of Ecosystem Functioning (CEFs), whose identification requires the collaboration of physical and chemical oceanography, biogeochemistry, marine geology, plankton, nekton and benthos ecology and biology, food web dynamics, marine biogeography. CEFs are fuzzy objects that reflect the instability of marine systems.

Mediterranean Bioconstructions Along the Italian Coast.

Marine bioconstructions are biodiversity-rich, three-dimensional biogenic structures, regulating key ecological functions of benthic ecosystems worldwide. Tropical coral reefs are outstanding for their beauty, diversity and complexity, but analogous types of bioconstructions are also present in temperate seas. The main bioconstructions in the Mediterranean Sea are represented by coralligenous formations, vermetid reefs, deep-sea cold-water corals, Lithophyllum byssoides trottoirs, coral banks formed by the shallow-water corals Cladocora caespitosa or Astroides calycularis, and sabellariid or serpulid worm reefs. Bioconstructions change the morphological and chemicophysical features of primary substrates and create new habitats for a large variety of organisms, playing pivotal roles in ecosystem functioning. In spite of their importance, Mediterranean bioconstructions have not received the same attention that tropical coral reefs have, and the knowledge of their biology, ecology and distribution is still fragmentary. All existing data about the spatial distribution of Italian bioconstructions have been collected, together with information about their growth patterns, dynamics and connectivity. The degradation of these habitats as a consequence of anthropogenic pressures (pollution, organic enrichment, fishery, coastal development, direct physical disturbance), climate change and the spread of invasive species was also investigated. The study of bioconstructions requires a holistic approach leading to a better understanding of their ecology and the application of more insightful management and conservation measures at basin scale, within ecologically coherent units based on connectivity: the cells of ecosystem functioning.

Correction: Successional dynamics of marine fouling hydroids (Cnidaria: Hydrozoa) at a finfish aquaculture facility in the Mediterranean Sea.

[This corrects the article DOI: 10.1371/journal.pone.0195352.].

A review of Paul Lassenius Kramp's hydrozoan types (taxonomic status, location and condition of the material) and a biography of the Danish zoologist.

During his scientific career, largely from 1910-1970, the Danish cnidarian specialist P.L. Kramp authored 94 scientific papers, and more than 30 other kinds of publications, such as reports and popular articles. Kramp was affiliated with the marine department of the Zoological Museum of the University of Copenhagen (now: The Natural History Museum of Denmark, NHMD), an institution intensely involved at the time in expeditions, in the assembling and maintenance of collections, and in the revision of a wide range of animal groups. Kramp took part in several expeditions in different parts of the world. He had opportunities to receive training in sampling and preservation techniques, to perform observations on living or at least freshly preserved material, and to collect specimens. Kramp dedicated most of his long career to studying the taxonomy of medusae, with some attention also to their polyp stages, describing 101 nominal hydrozoan taxa, including one order, four families and fifteen genera. Sixty-five of these taxa (one order, three families, twelve genera, 48 species and one subspecies) are still accepted in the original nominal form. Kramp raised the international reputation of his museum and his scientific results are still appreciated today.

Successional dynamics of marine fouling hydroids (Cnidaria: Hydrozoa) at a finfish aquaculture facility in the Mediterranean Sea.

Aquaculture is increasing rapidly to meet global seafood demand. Some hydroid populations have been linked to mortality and health issues in finfish and shellfish, but their dynamics in and around aquaculture farms remain understudied. In the present work, two experiments, each with 36 panels, tested colonization (factors: depth, season of immersion) and succession (factors: depth, submersion duration) over one year. Hydroid surface cover was estimated for each species, and data were analyzed with multivariate techniques. The assemblage of hydrozoans was species-poor, although species richness, frequency and abundance increased with time, paralleling the overall increase in structural complexity of fouling assemblages. Submersion duration and season of immersion were particularly important in determining the species composition of the assemblages in the succession and colonization experiments, respectively. Production of water-borne propagules, including medusae, from the hydroids was observed from locally abundant colonies, among them the well-known fouling species Obelia dichotoma, potentially representing a nuisance for cultured fish through contact-driven envenomations and gill disorders. The results illustrate the potential importance of fouling hydroids and their medusae to the health of organisms in the aquaculture industry.

Aglaophenia octodonta (Cnidaria, Hydrozoa) and the Associated Microbial Community: a Cooperative Alliance?

Recently, genetic approaches have revealed a surprising bacterial world as well as a growing knowledge of the enormous distribution of animal-bacterial interactions. In the present study, the diversity of the microorganisms associated to the hydroid Aglaophenia octodonta was studied with epifluorescence, optical, and scanning electron microscopy. Small subunit ribosomal RNA gene sequencing with "universal" and taxon-specific primers allowed the assignment of the microalgae to Symbiodinium and the peritrich ciliates to Pseudovorticella, while the luminous vibrios were identified as Vibrio jasicida of the Harvey clade. To understand the possible relationships among Vibrio jasicida, Symbiodinium, A. octodonta, and Pseudovorticella, specific treatments were conducted in microcosm experiments, with the antibiotic ampicillin and other substances that interfere with bacterial and hydroid metabolism. Treatment of A. octodonta with ampicillin resulted in a decrease of bacterial luminescence followed by Pseudovorticella detachment and Symbiodinium expulsion and suggesting that these microorganisms form a "consortium" with beneficial metabolic interdependence. This hypothesis was reinforced by the evidence that low concentrations of hydrogen peroxide, which stimulate the bacterial oxidative metabolism and luminescence by releasing oxygen, were able to counteract the detrimental effect of ampicillin on the stability of the studied A. octodonta association. A model is proposed in which microalgae that release oxygen during photosynthesis are useful to luminous bacteria for their metabolism and for establishing/maintaining symbiosis leading to a close alliance and mutual benefit of the system A. octodonta-Vibrio jasicida-Pseudovorticella sp.-Symbiodinium sp.

Deterministic Factors Overwhelm Stochastic Environmental Fluctuations as Drivers of Jellyfish Outbreaks.

Jellyfish outbreaks are increasingly viewed as a deterministic response to escalating levels of environmental degradation and climate extremes. However, a comprehensive understanding of the influence of deterministic drivers and stochastic environmental variations favouring population renewal processes has remained elusive. This study quantifies the deterministic and stochastic components of environmental change that lead to outbreaks of the jellyfish Pelagia noctiluca in the Mediterranen Sea. Using data of jellyfish abundance collected at 241 sites along the Catalan coast from 2007 to 2010 we: (1) tested hypotheses about the influence of time-varying and spatial predictors of jellyfish outbreaks; (2) evaluated the relative importance of stochastic vs. deterministic forcing of outbreaks through the environmental bootstrap method; and (3) quantified return times of extreme events. Outbreaks were common in May and June and less likely in other summer months, which resulted in a negative relationship between outbreaks and SST. Cross- and along-shore advection by geostrophic flow were important concentrating forces of jellyfish, but most outbreaks occurred in the proximity of two canyons in the northern part of the study area. This result supported the recent hypothesis that canyons can funnel P. noctiluca blooms towards shore during upwelling. This can be a general, yet unappreciated mechanism leading to outbreaks of holoplanktonic jellyfish species. The environmental bootstrap indicated that stochastic environmental fluctuations have negligible effects on return times of outbreaks. Our analysis emphasized the importance of deterministic processes leading to jellyfish outbreaks compared to the stochastic component of environmental variation. A better understanding of how environmental drivers affect demographic and population processes in jellyfish species will increase the ability to anticipate jellyfish outbreaks in the future.

Missing species among Mediterranean non-Siphonophoran Hydrozoa.

Hydrozoa of the Mediterranean Sea are well known and a recent monograph covers 457 species. Mediterranean non-Siphonophoran Hydrozoa comprises 398 species, an increasing number due to continuous updates, representing about 10 % of the 3,702 currently valid species reported in a recent world assessment of hydrozoan diversity. Many new records are non indigenous species, previously described species that occurred elsewhere and whose arrival was presumably caused by human activities. However, many species reported in the past are not recorded in recent times. Realistic assessments of species pools require addition of new species, but also subtraction of species not found since a certain period. With the confidence of extinction index, cases of putative extinction can be raised. Out of the 398 known species, only 162 (41 %) have been reported in the last decade, while 53 (13 %) are not recorded in the literature since at least 41 years. According to the confidence of extinction index, 60 % of the 53 missing species are extinct, and 11 % are putatively extinct from the basin. From a biogeographical point of view, the missing species are: 34 % endemic, 19 % boreal, 15 % Mediterranean-Atlantic, 11 % Indo-Pacific, 11 % circumtropical, 4 % cosmopolitan, 2 % tropical-Atlantic, 4 % non-classifiable. Fluctuations in species composition into a certain area cause heavy variability in the expression of both structural and functional biodiversity. As consequence, the regional biodiversity should be analyzed through its temporal evolution, to detect changes and their possible causes. This approach has profound consequences on biodiversity assessments and also on the compilation of red lists.

From Darwin's Origin of Species toward a theory of natural history.

Darwin is the father of evolutionary theory because he identified evolutionary patterns and, with Natural Selection, he ascertained the exquisitely ecological ultimate processes that lead to evolution. The proximate processes of evolution he proposed, however, predated the discovery of genetics, the backbone of modern evolutionary theory. The later discovery of the laws of inheritance by Mendel and the rediscovery of Mendel in the early 20th century led to two reforms of Darwinism: Neo-Darwinism and the Modern Synthesis (and subsequent refinements). If Darwin's evolutionary thought required much refinement, his ecological insight is still very modern. In the first edition of The Origin of Species, Darwin did not use either the word "evolution" or the word "ecology". "Ecology" was not coined until after the publication of the Origin. Evolution, for him, was the origin of varieties, then species, which he referred to as well-marked varieties, whereas, instead of using ecology, he used "the economy of nature". The Origin contains a high proportion of currently accepted ecological principles. Darwin labelled himself a naturalist. His discipline (natural history) was a blend of ecology and evolution in which he investigated both the patterns and the processes that determine the organization of life. Reductionist approaches, however, often keep the two disciplines separated from each other, undermining a full understanding of natural phenomena that might be favored by blending ecology and evolution through the development of a modern Theory of Natural History based on Darwin's vision of the study of life.

The non-Siphonophoran Hydrozoa (Cnidaria) of Salento, Italy with notes on their life-cycles: an illustrated guide.

The majority of Hydrozoa is represented by not readily noticeable, small species. In recent decades, however, taxonomic knowledge of the group has increased worldwide, with a significant number of investigations focused on the Mediterranean Sea. Over more than two decades, 115 species of hydrozoans were recorded from coastal waters along nearly 300 km of the Salento Peninsula (Apulia, Italy). For each species, records from different collections were merged into single sheets of a general database. For each species, the following information is reported: description, cnidome, biology, occurrence in Salento, worldwide distribution, and bibliography. Descriptions refer to the benthic hydroid stage and, when present, also to the planktonic medusa stage. The 115 species of Hydrozoa, recorded along the Salento coastline, represent 25% of the Mediterranean Hydrozoa fauna (totaling 461 species), and nearly 3% of 3,702 world's known species covered in a recent monograph. Four species are non-indigenous, three of them with invasive behavior (Clytia hummelincki, Clytia linearis, and Eudendrium carneum), and one species now very common (Eudendrium merulum) in Salento. The complete life cycle of Clytia paulensis (Vanhöffen, 1910) is described for the first time.

Global warming and mass mortalities of benthic invertebrates in the Mediterranean Sea.

Satellite data show a steady increase, in the last decades, of the surface temperature (upper few millimetres of the water surface) of the Mediterranean Sea. Reports of mass mortalities of benthic marine invertebrates increased in the same period. Some local studies interpreted the two phenomena in a cause-effect fashion. However, a basin-wide picture of temperature changes combined with a systematic assessment on invertebrate mass mortalities was still lacking. Both the thermal structure of the water column in the Mediterranean Sea over the period 1945-2011 and all documented invertebrate mass mortality events in the basin are analysed to ascertain if: 1- documented mass mortalities occurred under conditions of positive temperature trends at basin scale, and 2- atypical thermal conditions were registered at the smaller spatial and temporal scale of mass mortality events. The thermal structure of the shallow water column over the last 67 years was reconstructed using data from three public sources: MEDAR-MEDATLAS, World Ocean Database, MFS-VOS programme. A review of the mass mortality events of benthic invertebrates at Mediterranean scale was also carried out. The analysis of in situ temperature profiles shows that the Mediterranean Sea changed in a non-homogeneous fashion. The frequency of mass mortalities is increasing. The areas subjected to these events correspond to positive thermal anomalies. Statistically significant temperature trends in the upper layers of the Mediterranean Sea show an increase of up to 0.07°C/yr for a large fraction of the basin. Mass mortalities are consistent with both the temperature increase at basin scale and the thermal changes at local scale, up to 5.2°C. Our research supports the existence of a causal link between positive thermal anomalies and observed invertebrate mass mortalities in the Mediterranean Sea, invoking focused mitigation initiatives in sensitive areas.

Predator decline leads to decreased stability in a coastal fish community.

Fisheries exploitation has caused widespread declines in marine predators. Theory predicts that predator depletion will destabilise lower trophic levels, making natural communities more vulnerable to environmental perturbations. However, empirical evidence has been limited. Using a community matrix model, we empirically assessed trends in the stability of a multispecies coastal fish community over the course of predator depletion. Three indices of community stability (resistance, resilience and reactivity) revealed significantly decreasing stability concurrent with declining predator abundance. The trophically downgraded community exhibited weaker top-down control, leading to predator-release processes in lower trophic levels and increased susceptibility to perturbation. At the community level, our results suggest that high predator abundance acts as a stabilising force to the naturally stochastic and highly autocorrelated dynamics in low trophic species. These findings have important implications for the conservation and management of predators in marine ecosystems and provide empirical support for the theory of predatory control.