Carrion ecology in inland aquatic ecosystems: a systematic review.

Carrion ecology, i.e. the decomposition and recycling of dead animals, has traditionally been neglected as a key process in ecosystem functioning. Similarly, despite the large threats that inland aquatic ecosystems (hereafter, aquatic ecosystems) face, the scientific literature is still largely biased towards terrestrial ecosystems. However, there has been an increasing number of studies on carrion ecology in aquatic ecosystems in the last two decades, highlighting their key role in nutrient recirculation and disease control. Thus, a global assessment of the ecological role of scavengers and carrion in aquatic ecosystems is timely. Here, we systematically reviewed scientific articles on carrion ecology in aquatic ecosystems to describe current knowledge, identify research gaps, and promote future studies that will deepen our understanding in this field. We found 206 relevant studies, which were highly biased towards North America, especially in lotic ecosystems, covering short time periods, and overlooking seasonality, a crucial factor in scavenging dynamics. Despite the low number of studies on scavenger assemblages, we recorded 55 orders of invertebrates from 179 families, with Diptera and Coleoptera being the most frequent orders. For vertebrates, we recorded 114 species from 40 families, with birds and mammals being the most common. Our results emphasise the significance of scavengers in stabilising food webs and facilitating nutrient cycling within aquatic ecosystems. Studies were strongly biased towards the assessment of the ecosystem effects of carrion, particularly of salmon carcasses in North America. The second most common research topic was the foraging ecology of vertebrates, which was mostly evaluated through sporadic observations of carrion in the diet. Articles assessing scavenger assemblages were scarce, and only a limited number of these studies evaluated carrion consumption patterns, which serve as a proxy for the role of scavengers in the ecosystem. The ecological functions performed by carrion and scavengers in aquatic ecosystems were diverse. The main ecological functions were carrion as food source and the role of scavengers in nutrient cycling, which appeared in 52.4% (N = 108) and 46.1% (N = 95) of publications, respectively. Ecosystem threats associated with carrion ecology were also identified, the most common being water eutrophication and carrion as source of pathogens (2.4%; N = 5 each). Regarding the effects of carrion on ecosystems, we found studies spanning all ecosystem components (N = 85), from soil or the water column to terrestrial vertebrates, with a particular focus on aquatic invertebrates and fish. Most of these articles found positive effects of carrion on ecosystems (e.g. higher species richness, abundance or fitness; 84.7%; N = 72), while a minority found negative effects, changes in community composition, or even no effects. Enhancing our understanding of scavengers and carrion in aquatic ecosystems is crucial to assessing their current and future roles amidst global change, mainly for water-land nutrient transport, due to changes in the amount and speed of nutrient movement, and for disease control and impact mitigation, due to the predicted increase in occurrence and magnitude of mortality events in aquatic ecosystems.

Role of climatic variability in shaping intraspecific variation of thermal tolerance in Mediterranean water beetles.

The climatic variability hypothesis (CVH) predicts that organisms in more thermally variable environments have wider thermal breadths and higher thermal plasticity than those from more stable environments. However, due to evolutionary trade-offs, taxa with greater absolute thermal limits may have little plasticity of such limits (trade-off hypothesis). The CVH assumes that climatic variability is the ultimate driver of thermal tolerance variation across latitudinal and altitudinal gradients, but average temperature also varies along such gradients. We explored intraspecific variation of thermal tolerance in three typical Mediterranean saline water beetles (families Hydrophilidae and Dytiscidae). For each species, we compared two populations where the species coexist, with similar annual mean temperature but contrasting thermal variability (continental vs. coastal population). We estimated thermal limits of adults from each population, previously acclimated at 17, 20, or 25 °C. We found species-specific patterns but overall, our results agree with the CVH regarding thermal ranges, which were wider in the continental (more variable) population. In the two hydrophilid species, this came at the cost of losing plasticity of the upper thermal limit in this population, supporting the trade-off hypothesis, but not in the dytiscid one. Our results support the role of local adaptation to thermal variability and trade-offs between basal tolerance and physiological plasticity in shaping thermal tolerance in aquatic ectotherms, but also suggest that intraspecific variation of thermal tolerance does not fit a general pattern among aquatic insects. Overlooking such intraspecific variation could lead to inaccurate predictions of the vulnerability of aquatic insects to global warming.

A horizon scan exercise for aquatic invasive alien species in Iberian inland waters.

As the number of introduced species keeps increasing unabatedly, identifying and prioritising current and potential Invasive Alien Species (IAS) has become essential to manage them. Horizon Scanning (HS), defined as an exploration of potential threats, is considered a fundamental component of IAS management. By combining scientific knowledge on taxa with expert opinion, we identified the most relevant aquatic IAS in the Iberian Peninsula, i.e., those with the greatest geographic extent (or probability of introduction), severe ecological, economic and human health impacts, greatest difficulty and acceptability of management. We highlighted the 126 most relevant IAS already present in Iberian inland waters (i.e., Concern list) and 89 with a high probability of being introduced in the near future (i.e., Alert list), of which 24 and 10 IAS, respectively, were considered as a management priority after receiving the highest scores in the expert assessment (i.e., top-ranked IAS). In both lists, aquatic IAS belonging to the four thematic groups (plants, freshwater invertebrates, estuarine invertebrates, and vertebrates) were identified as having been introduced through various pathways from different regions of the world and classified according to their main functional feeding groups. Also, the latest update of the list of IAS of Union concern pursuant to Regulation (EU) No 1143/2014 includes only 12 top-ranked IAS identified for the Iberian Peninsula, while the national lists incorporate the vast majority of them. This fact underlines the great importance of taxa prioritisation exercises at biogeographical scales as a step prior to risk analyses and their inclusion in national lists. This HS provides a robust assessment and a cost-effective strategy for decision-makers and stakeholders to prioritise the use of limited resources for IAS prevention and management. Although applied at a transnational level in a European biodiversity hotspot, this approach is designed for potential application at any geographical or administrative scale, including the continental one.

Towards evidence-based conservation of subterranean ecosystems.

Subterranean ecosystems are among the most widespread environments on Earth, yet we still have poor knowledge of their biodiversity. To raise awareness of subterranean ecosystems, the essential services they provide, and their unique conservation challenges, 2021 and 2022 were designated International Years of Caves and Karst. As these ecosystems have traditionally been overlooked in global conservation agendas and multilateral agreements, a quantitative assessment of solution-based approaches to safeguard subterranean biota and associated habitats is timely. This assessment allows researchers and practitioners to understand the progress made and research needs in subterranean ecology and management. We conducted a systematic review of peer-reviewed and grey literature focused on subterranean ecosystems globally (terrestrial, freshwater, and saltwater systems), to quantify the available evidence-base for the effectiveness of conservation interventions. We selected 708 publications from the years 1964 to 2021 that discussed, recommended, or implemented 1,954 conservation interventions in subterranean ecosystems. We noted a steep increase in the number of studies from the 2000s while, surprisingly, the proportion of studies quantifying the impact of conservation interventions has steadily and significantly decreased in recent years. The effectiveness of 31% of conservation interventions has been tested statistically. We further highlight that 64% of the reported research occurred in the Palearctic and Nearctic biogeographic regions. Assessments of the effectiveness of conservation interventions were heavily biased towards indirect measures (monitoring and risk assessment), a limited sample of organisms (mostly arthropods and bats), and more accessible systems (terrestrial caves). Our results indicate that most conservation science in the field of subterranean biology does not apply a rigorous quantitative approach, resulting in sparse evidence for the effectiveness of interventions. This raises the important question of how to make conservation efforts more feasible to implement, cost-effective, and long-lasting. Although there is no single remedy, we propose a suite of potential solutions to focus our efforts better towards increasing statistical testing and stress the importance of standardising study reporting to facilitate meta-analytical exercises. We also provide a database summarising the available literature, which will help to build quantitative knowledge about interventions likely to yield the greatest impacts depending upon the subterranean species and habitats of interest. We view this as a starting point to shift away from the widespread tendency of recommending conservation interventions based on anecdotal and expert-based information rather than scientific evidence, without quantitatively testing their effectiveness.

Climatic stability, not average habitat temperature, determines thermal tolerance of subterranean beetles.

The climatic variability hypothesis predicts the evolution of species with wide thermal tolerance ranges in environments with variable temperatures, and the evolution of thermal specialists in thermally stable environments. In caves, the extent of spatial and temporal thermal variability experienced by taxa decreases with their degree of specialization to deep subterranean habitats. We use phylogenetic generalized least squares to model the relationship among thermal tolerance (upper lethal limits), subterranean specialization (estimated using ecomorphological traits), and habitat temperature in 16 beetle species of the tribe Leptodirini (Leiodidae). We found a significant, negative relationship between thermal tolerance and the degree of subterranean specialization. Conversely, habitat temperature had only a marginal effect on lethal limits. In agreement with the climatic variability hypothesis and under a climate change context, we show that the specialization process to live in deep subterranean habitats involves a reduction of upper lethal limits, but not an adjustment to habitat temperature. Thermal variability seems to exert a higher evolutionary pressure than mean habitat temperature to configure the thermal niche of subterranean species. Our results provide novel insights on thermal physiology of species with poor dispersal capabilities and on the evolutionary process of adaptation to subterranean environments. We further emphasize that the pathways determining vulnerability of subterranean species to climate change greatly depend on the degree of specialization to deep subterranean environments.

Beyond survival experiments: using biomarkers of oxidative stress and neurotoxicity to assess vulnerability of subterranean fauna to climate change.

Accurate assessments of species vulnerability to climate change need to consider the physiological capacity of organisms to deal with temperature changes and identify early signs of thermally induced stress. Oxidative stress biomarkers and acetylcholinesterase activity are useful proxies of stress at the cellular and nervous system level. Such responses are especially relevant for poor dispersal organisms with limited capacity for behavioural thermoregulation, like deep subterranean species. We combined experimental measurements of upper lethal thermal limits, acclimation capacity and biomarkers of oxidative stress and neurotoxicity to assess the impact of heat stress (20°C) at different exposure times (2 and 7 days) on the Iberian endemic subterranean beetle Parvospeonomus canyellesi. Survival response (7 days of exposure) was similar to that reported for other subterranean specialist beetles (high survival up to 20°C but no above 23°C). However, a low physiological plasticity (i.e. incapacity to increase heat tolerance via acclimation) and signs of impairment at the cellular and nervous system level were observed after 7 days of exposure at 20°C. Such sublethal effects were identified by significant differences in total antioxidant capacity, glutathione S-transferase activity, the ratio of reduced to oxidized forms of glutathione and acetylcholinesterase activity between the control (cave temperature) and 20°C treatment. At 2 days of exposure, most biomarker values indicated some degree of oxidative stress in both the control and high-temperature treatment, likely reflecting an initial altered physiological status associated to factors other than temperature. Considering these integrated responses and the predicted increase in temperature in its unique locality, P. canyellesi would have a narrower thermal safety margin to face climate change than that obtained considering only survival experiments. Our results highlight the importance of exploring thermally sensitive processes at different levels of biological organization to obtain more accurate estimates of the species capacity to face climate change.

Heat tolerance and acclimation capacity in subterranean arthropods living under common and stable thermal conditions.

Cave-dwelling ectotherms, which have evolved for millions of years under stable thermal conditions, could be expected to have adjusted their physiological limits to the narrow range of temperatures they experience and to be highly vulnerable to global warming. However, most of the few existing studies on thermal tolerance in subterranean invertebrates highlight that despite the fact that they show lower heat tolerance than most surface-dwelling species, their upper thermal limits are generally not adjusted to ambient temperature. The question remains to what extent this pattern is common across subterranean invertebrates. We studied basal heat tolerance and its plasticity in four species of distant arthropod groups (Coleoptera, Diplopoda, and Collembola) with different evolutionary histories but under similar selection pressures, as they have been exposed to the same constant environmental conditions for a long time. Adults were exposed at different temperatures for 1 week to determine upper lethal temperatures. Then, individuals from previous sublethal treatments were transferred to a higher temperature to determine acclimation capacity. Upper lethal temperatures of three of the studied species were similar to those reported for other subterranean species (between 20 and 25°C) and widely exceeded the cave temperature (13-14°C). The diplopod species showed the highest long-term heat tolerance detected so far for a troglobiont (i.e., obligate subterranean) species (median lethal temperature after 7 days exposure: 28°C) and a positive acclimation response. Our results agree with previous studies showing that heat tolerance in subterranean species is not determined by environmental conditions. Thus, subterranean species, even those living under similar climatic conditions, might be differently affected by global warming.

Do all roads lead to Rome? Exploring community trajectories in response to anthropogenic salinization and dilution of rivers.

Abiotic stress shapes how communities assemble and support ecological functions. However, it remains unclear whether artificially increasing or decreasing stress levels would lead to communities assembling predictably along a single axis of variation or along multiple context-dependent trajectories of change. In response to stress intensity alterations, we hypothesize that a single trajectory of change occurs when trait-based assembly prevails, while multiple trajectories of change arise when dispersal-related processes modify colonization and trait-filtering dynamics. Here, we tested these hypotheses using aquatic macroinvertebrates from rivers exposed to gradients of natural salinity and artificially diluted or salinized ion contents. Our results showed that trait-filtering was important in driving community assembly in natural and diluted rivers, while dispersal-related processes seemed to play a relevant role in response to salinization. Salinized rivers showed novel communities with different trait composition, while natural and diluted communities exhibited similar taxonomic and trait compositional patterns along the conductivity gradient. Our findings suggest that the artificial modification of chemical stressors can result in different biological communities, depending on the direction of the change (salinization or dilution), with trait-filtering, and organism dispersal and colonization dynamics having differential roles in community assembly. The approach presented here provides both empirical and conceptual insights that can help in anticipating the ecological effects of global change, especially for those stressors with both natural and anthropogenic origins.This article is part of the theme issue 'Salt in freshwaters: causes, ecological consequences and future prospects'.

Insect communities in saline waters consist of realized but not fundamental niche specialists.

Considering how organisms adapt to stress is essential if we are to anticipate biological responses to global change in ecosystems. Communities in stressful environments can potentially be assembled by specialists (i.e. species that only occur in a limited range of environmental conditions) and/or generalist species with wider environmental tolerances. We review the existing literature on the salinity tolerance of aquatic insects previously identified as saline specialists because they were exclusively found in saline habitats, and explore if these saline realized niche specialists are also specialists in their fundamental niches or on the contrary are fundamental niche generalist species confined to the highest salinities they can tolerate. The results suggest that species inhabiting saline waters are generalists in their fundamental niches, with a predominant pattern of high survival in freshwater-low salinity conditions, where their fitness tends to be similar or even higher than in saline waters. Additionally, their performance in freshwater tends to be similar to related strictly freshwater species, so no apparent trade-off of generalization is shown. These results are discussed in the framework of the ecological and evolutionary processes driving community assembly across the osmotic stress gradient, and their potential implications for predicting impacts from saline dilution and freshwater salinization.This article is part of the theme issue 'Salt in freshwaters: causes, ecological consequences and future prospects'.

Effects of salinity changes on aquatic organisms in a multiple stressor context.

Under global change, the ion concentration of aquatic ecosystems is changing worldwide. Many freshwater ecosystems are being salinized by anthropogenic salt inputs, whereas many naturally saline ones are being diluted by agricultural drainages. This occurs concomitantly with changes in other stressors, which can result in additive, antagonistic or synergistic effects on organisms. We reviewed experimental studies that manipulated salinity and other abiotic stressors, on inland and transitional aquatic habitats, to (i) synthesize their main effects on organisms' performance, (ii) quantify the frequency of joint effect types across studies and (iii) determine the overall individual and joint effects and their variation among salinity-stressor pairs and organism groups using meta-analyses. Additive effects were slightly more frequent (54%) than non-additive ones (46%) across all the studies (n = 105 responses). However, antagonistic effects were dominant for the stressor pair salinity and toxicants (44%, n = 43), transitional habitats (48%, n = 31) and vertebrates (71%, n = 21). Meta-analyses showed detrimental additive joint effects of salinity and other stressors on organism performance and a greater individual impact of salinity than the other stressors. These results were consistent across stressor pairs and organism types. These findings suggest that strategies to mitigate multiple stressor impacts on aquatic ecosystems should prioritize restoring natural salinity concentrations.This article is part of the theme issue 'Salt in freshwaters: causes, ecological consequences and future prospects'.

Environmental niche unfilling but limited options for range expansion by active dispersion in an alien cavity-nesting wasp.

BACKGROUND: Predicting the patterns of range expansion of alien species is central to develop effective strategies for managing potential biological invasions. Here, we present a study on the potential distribution of the American cavity-nesting, Orthoptera-hunting and solitary wasp, Isodontia mexicana (Hymenoptera: Sphecidae), which was first detected as alien species in France in 1960 and now is present in many European countries. After having updated its current distribution, we estimated the environmental space (based on bioclimatic data and altitude) occupied by the species and subsequently predicted its environmental potential distribution under both present and future climatic conditions at global scale. RESULTS: The wasp lives in low-altitude areas of the Northern hemisphere with moderate temperatures and precipitation. The environmental space occupied in the invaded area is practically just a subset (42%) of that occupied in the native area, showing a process of environmental niche unfilling (i.e. the species only partially fills its environmental niche in the invaded range). Besides, I. mexicana could also live in other temperate areas, mainly in the Southern hemisphere, particularly close to the coasts. However, geographic (oceans) and/or climatic (tropical areas, mountain chains) barriers would prevent the species to reach these potential areas unless through human trade activity. The species could thus only reach, by active dispersion, the remaining invadable areas of Europe. Estimations for the future (2050 and 2070) predict an expansion through active dispersion towards North in the native range and towards North and East in the invaded range, but future conditions would not break down the current climatic barriers in the Southern hemisphere. CONCLUSIONS: Isodontia mexicana has not shifted its environmental niche in the invaded area. It could still occupy some new areas by active dispersion, but confined to Europe. The conspicuous niche unfilling shown by this wasp species could reflect the likely single introduction in Europe just a few decades ago. Furthermore, results stay in line with other studies that found niche unfilling rather than niche expansion in insects.

Vulnerabilidad de las especies de Dytiscidae (Coleoptera) en Cuba / Vulnerability of Dytiscidae species (Coleoptera) in Cuba

Resumen La diversidad de coleópteros acuáticos de Cuba se encuentra entre las más elevadas en el Caribe insular. Sin embargo, existe escasa información sobre los ditíscidos cubanos para identificar las especies y hábitats que requieren urgentes acciones de conservación. El objetivo de este estudio fue identificar los taxa amenazados de la familia Dytiscidae en Cuba según su grado de vulnerabilidad. Para ello se recopiló información sobre la distribución de todos los ditíscidos cubanos, incluyendo datos de la literatura, de colecciones y de muestreos realizados en el período comprendido entre los años 2000 y 2014. Además, fue evaluada la vulnerabilidad de las especies a partir de una metodología que combina seis criterios referentes a las características de las especies y los hábitats que ocupan. El análisis de vulnerabilidad fue posible realizarlo a 50 de las 53 especies de ditíscidos (94 %) presentes en Cuba. Un total de cinco especies (9 %) presentan vulnerabilidad alta, 20 (38 %) media y 25 (48 %) baja. Las cinco especies más amenazadas son Laccodytes cobrinae, Laccophilus alariei, Copelatus barbouri, C. darlingtoni, and Desmopachria glabella. Estas son endémicas de Cuba y se caracterizan por presentar poblaciones muy localizadas, poco abundantes, así como por una alta especificidad de hábitat. Por ello, se propone su inclusión en lista roja de la fauna de Cuba, así como en la Lista Roja de la UICN bajo la categoría de "Vulnerable". Además, no todas las áreas donde se encuentran estas especies pertenecen al Sistema Nacional de Áreas Protegidas de Cuba, por lo que recomendamos la inclusión de la Laguna Base Julio A. Mella y el Norte de Imías en este sistema. Así mismo, sugerimos la implementación de medidas de gestión efectivas que eviten o mitiguen la alteración de los ecosistemas acuáticos en los que aparecen las especies vulnerables.

Abstract Cuba has one of the richest diving beetles species diversity in the Caribbean islands. However, Cuban Dytiscidae remain scarcely studied, and there is need to identify those species and habitats that urgently require effective conservation actions. Here we aim to identify the threatened taxa of the family Dytiscidae in Cuba according to their degree of vulnerability. For that, we compiled distributional data on the Cuban fauna, including data from literature, collections and own samplings carried out between the period 2000 and 2014. In addition, the vulnerability of the species was evaluated using a methodology that combines six criteria regarding both species and habitat attributes. This analysis of vulnerability was completed to 50 (94 %) out of 53 Cuban species. Five species (9 %) were identified as highly vulnerable, 20 (38 %) moderately and 25 (48 %) as having low conservation status. The five most threatened species were Laccodytes cobrinae, Laccophilus alariei, Copelatus barbouri, C. darlingtoni, and Desmopachria glabella. These species are endemic to Cuba and characterized by highly localized populations and low abundance as well as high habitat specificity. Thus, we propose the inclusion of these five species in both the red list of Cuban fauna and the IUCN Red List (under the category of "Vulnerable"). In addition, not all areas where these species were found belong to the National System of Protected Areas of Cuba. Thus, we suggest the inclusion of the Laguna Base Julio A. Mella and the North of Imías, and to develop effective management measures to prevent the alteration of these aquatic ecosystems. Rev. Biol. Trop. 66(2): 709-721. Epub 2018 June 01.

Matches and mismatches between conservation investments and biodiversity values in the European Union.

Recently, the European Commission adopted a new strategy to halt the loss of biodiversity. Member states are expected to favor a more effective collection and redistribution of European Union (EU) funds under the current Multiannual Financial Framework for 2014-2020. Because of the large spatial variation in the distribution of biodiversity and conservation needs at the continental scale, EU instruments should ensure that countries with higher biodiversity values get more funds and resources for the conservation than other countries. Using linear regressions, we assessed the association between conservation investments and biodiversity values across member states, accounting for a variety of conservation investment indicators, taxonomic groups (including groups of plants, vertebrates, and invertebrates), and indicators of biodiversity value. In general, we found clear overall associations between conservation investments and biodiversity variables. However, some countries received more or less investment than would be expected based on biodiversity values in those countries. We also found that the extensive use of birds as unique indicators of conservation effectiveness may lead to biased decisions. Our results can inform future decisions regarding funding allocation and thus improve distribution of EU conservation funds.

Thermal niche estimators and the capability of poor dispersal species to cope with climate change.

For management strategies in the context of global warming, accurate predictions of species response are mandatory. However, to date most predictions are based on niche (bioclimatic) models that usually overlook biotic interactions, behavioral adjustments or adaptive evolution, and assume that species can disperse freely without constraints. The deep subterranean environment minimises these uncertainties, as it is simple, homogeneous and with constant environmental conditions. It is thus an ideal model system to study the effect of global change in species with poor dispersal capabilities. We assess the potential fate of a lineage of troglobitic beetles under global change predictions using different approaches to estimate their thermal niche: bioclimatic models, rates of thermal niche change estimated from a molecular phylogeny, and data from physiological studies. Using bioclimatic models, at most 60% of the species were predicted to have suitable conditions in 2080. Considering the rates of thermal niche change did not improve this prediction. However, physiological data suggest that subterranean species have a broad thermal tolerance, allowing them to stand temperatures never experienced through their evolutionary history. These results stress the need of experimental approaches to assess the capability of poor dispersal species to cope with temperatures outside those they currently experience.

Similarity in the difference: changes in community functional features along natural and anthropogenic stress gradients.

The effect of stressors on biodiversity can vary in relation to the degree to which biological communities have adapted over evolutionary time. We compared the responses of functional features of stream insect communities along chronic stress gradients with contrasting time persistence. Water salinity and land use intensification were used as examples of natural (long-term persistent) and anthropogenic (short-term persistent) stressors, respectively. A new trait-based approach was applied to quantify functional diversity components and functional redundancy within the same multidimensional space, using metrics at the taxon and community levels. We found similar functional responses along natural and anthropogenic stress gradients. In both cases, the mean taxon functional richness and functional similarity between taxa increased with stress, whereas community functional richness and functional redundancy decreased. Despite the differences in evolutionary persistence, both chronic stressors act as strong nonrandom environmental filters, producing convergent functional responses. These results can improve our ability to predict functional effects of novel stressors at ecoloiical and evolutionary scales.

Atlas of Iberian water beetles (ESACIB database).

The ESACIB ('EScarabajos ACuáticos IBéricos') database is provided, including all available distributional data of Iberian and Balearic water beetles from the literature up to 2013, as well as from museum and private collections, PhD theses, and other unpublished sources. The database contains 62,015 records with associated geographic data (10×10 km UTM squares) for 488 species and subspecies of water beetles, 120 of them endemic to the Iberian Peninsula and eight to the Balearic Islands. This database was used for the elaboration of the "Atlas de los Coleópteros Acuáticos de España Peninsular". In this dataset data of 15 additional species has been added: 11 that occur in the Balearic Islands or mainland Portugal but not in peninsular Spain and an other four with mainly terrestrial habits within the genus Helophorus (for taxonomic coherence). The complete dataset is provided in Darwin Core Archive format.

Lack of evolutionary adjustment to ambient temperature in highly specialized cave beetles.

BACKGROUND: A key question in evolutionary biology is the relationship between species traits and their habitats. Caves offer an ideal model to test the adjustment of species to their surrounding temperature, as they provide homogeneous and simple environments. We compared two species living under different thermal conditions within a lineage of Pyrenean beetles highly modified for the subterranean life since the Miocene. One, Troglocharinus fonti, is found in caves at 4-11°C in the ancestral Pyrenean range. The second, T. ferreri, inhabits the coastal area of Catalonia since the early Pliocene, and lives at 14-16°C. RESULTS: We found no differences in their short term upper thermal limit (ca. 50°C), similar to that of most organisms, or their lower thermal limit (ca. -2.5°C), higher than for most temperate insects and suggesting the absence of cryoprotectants. In longer term tests (7 days) survival between 6-20°C was almost 100% for both species plus two outgroups of the same lineage, but all four died between 23-25°C, without significant differences between them. CONCLUSIONS: Our results suggest that species in this lineage have lost some of the thermoregulatory mechanisms common in temperate insects, as their inferred default tolerance range is larger than the thermal variation experienced through their whole evolutionary history.

Thermal niche evolution and geographical range expansion in a species complex of western Mediterranean diving beetles.

BACKGROUND: Species thermal requirements are one of the principal determinants of their ecology and biogeography, although our understanding of the interplay between these factors is limited by the paucity of integrative empirical studies. Here we use empirically collected thermal tolerance data in combination with molecular phylogenetics/phylogeography and ecological niche modelling to study the evolution of a clade of three western Mediterranean diving beetles, the Agabus brunneus complex. RESULTS: The preferred mitochondrial DNA topology recovered A. ramblae (North Africa, east Iberia and Balearic islands) as paraphyletic, with A. brunneus (widespread in the southwestern Mediterranean) and A. rufulus (Corsica and Sardinia) nested within it, with an estimated origin between 0.60-0.25 Ma. All three species were, however, recovered as monophyletic using nuclear DNA markers. A Bayesian skyline plot suggested demographic expansion in the clade at the onset of the last glacial cycle. The species thermal tolerances differ significantly, with A. brunneus able to tolerate lower temperatures than the other taxa. The climatic niche of the three species also differs, with A. ramblae occupying more arid and seasonal areas, with a higher minimum temperature in the coldest month. The estimated potential distribution for both A. brunneus and A. ramblae was most restricted in the last interglacial, becoming increasingly wider through the last glacial and the Holocene. CONCLUSIONS: The A. brunneus complex diversified in the late Pleistocene, most likely in south Iberia after colonization from Morocco. Insular forms did not differentiate substantially in morphology or ecology, but A. brunneus evolved a wider tolerance to cold, which appeared to have facilitated its geographic expansion. Both A. brunneus and A. ramblae expanded their ranges during the last glacial, although they have not occupied areas beyond their LGM potential distribution except for isolated populations of A. brunneus in France and England. On the islands and possibly Tunisia secondary contact between A. brunneus and A. ramblae or A. rufulus has resulted in introgression. Our work highlights the complex dynamics of speciation and range expansions within southern areas during the last glacial cycle, and points to the often neglected role of North Africa as a source of European biodiversity.

A new species of Desmopachria Babington (Coleoptera: Dytiscidae) from Cuba with a prediction of its geographic distribution and notes on other Cuban species of the genus.

A new species, Desmopachria andreae sp. n. is described from Cuba. Diagnostic characters including illustrations of male genitalia are provided and illustrated for the five species of the genus occurring on the island. For these five species both a simple key to adults and maps of their known distribution in Cuba are also provided. Using a Maximun Entropy method (MaxEnt), a distribution model was developed for D. andreae sp.n. Based on the model's predictions, this species has a higher probability of occurring in high altitude forests (above 1000 m a.s.l.), characterised by relatively low temperatures especially during the hottest and wettest seasons, specifically, the mountainous areas of the Macizo de Guamuhaya (Central Cuba), Sierra Maestra (S Cuba) and Nipe-Sagua-Baracoa (NE Cuba). In some of these areas the species has not yet been recorded, and should be searched for in future field surveys.