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An intricate interplay between evolutionary and demographic processes has frequently resulted in complex patterns of genetic and phenotypic diversity in alpine lineages, posing serious challenges to species delimitation and biodiversity conservation planning. Here we integrate genomic data, geometric morphometric analyses and thermal tolerance experiments to explore the role of Pleistocene climatic changes and adaptation to alpine environments on patterns of genomic and phenotypic variation in diving beetles from the taxonomically complex Agabus bipustulatus species group. Genetic structure and phylogenomic analyses revealed the presence of three geographically cohesive lineages, two representing trans-Palearctic and Iberian populations of the elevation-generalist A. bipustulatus and another corresponding to the strictly-alpine A. nevadensis, a narrow-range endemic taxon from the Sierra Nevada mountain range in southeastern Iberia. The best-supported model of lineage divergence, along with the existence of pervasive genetic introgression and admixture in secondary contact zones, is consistent with a scenario of population isolation and connectivity linked to Quaternary climatic oscillations. Our results suggest that A. nevadensis is an alpine ecotype of A. bipustulatus, whose genotypic, morphological and physiological differentiation likely resulted from an interplay between population isolation and local altitudinal adaptation. Remarkably, within the Iberian Peninsula, such ecotypic differentiation is unique to Sierra Nevada populations and has not been replicated in other alpine populations of A. bipustulatus. Collectively, our study supports fast ecotypic differentiation and incipient speciation processes within the study complex and points to Pleistocene glaciations and local adaptation along elevational gradients as key drivers of biodiversity generation in alpine environments.
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Besouros , Especiação Genética , Genética Populacional , Filogenia , Animais , Besouros/genética , Besouros/classificação , Besouros/anatomia & histologia , Ecótipo , Fenótipo , Espanha , Genótipo , Variação GenéticaRESUMO
BACKGROUND: Although the important roles of proprioception and neuromuscular control in carpal instabilities under laboratory conditions have been recognized, only a few studies have translated this knowledge into a routine clinical practice. PURPOSE: This study aimed to evaluate the results of a personalized rehabilitation in patients with carpal instability on functionality and pain intensity. STUDY DESIGN: This was a case series study. METHODS: This case series included 39 adults (mean age: 38.2 ± 14.0 years; 16/23 females/males) diagnosed with carpal instability (radial or ulnar) with indication for orthopedic treatment. The disabilities of the arm, shoulder, and hand questionnaire was used to assess upper limb functionality. Pain perception was assessed using a visual analog scale. Exercise-based physiotherapy interventions were performed according to the clinical needs of the patients for at least 6 weeks (2-3 sessions per week). For the treatment of radial instability (n = 13), strengthening exercises of the abductor pollicis longus, extensor carpi radialis longus, flexor carpi radialis, and pronator quadratus muscles were prescribed. For the treatment of ulnar instability (n = 24), extensor carpi ulnaris and pronator quadratus were trained. All patients underwent proprioceptive training in open kinetic chain and closed kinetic chain, as well as strengthening of the unaffected hand. Changes before and after treatment were compared using the nonparametric Wilcoxon signed rank test. RESULTS: A significant improvement with a large effect size in disabilities of the arm, shoulder, and hand (P < .001; d = 2.9) and visual analog scale (P < .001; d = 3.2) scores were obtained after treatment. Moreover, the changes were greater than the minimal clinically important difference of 10.8 and 1.4, respectively. Similar results were found when patients with radial instability and ulnar instability were analyzed separately. CONCLUSIONS: Personalized training with specific proprioception and strengthening exercises produces improvements in functionality and pain perception in our cohort of people with carpal instability. These results highlight the importance of multicomponent exercise in the treatment of wrist instability. Future randomized clinical trials should further investigate the effectiveness of this protocol.
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The study of the genetic makeup and demographic fate of alien species is essential to understand their capacity to recover from founder effects, adapt to new environmental conditions and, ultimately, become invasive and potentially damaging. Here, we employ genomic data to gain insights into key demographic processes that might help to explain the extraordinarily successful invasion of the Western Mediterranean region by the North American boatman Trichocorixa verticalis (Hemiptera: Corixidae). Our analyses revealed the genetic distinctiveness of populations from the main areas comprising the invasive range and coalescent-based simulations supported that they originated from independent introductions events probably involving different source populations. Testing of alternative demographic models indicated that all populations experienced a strong bottleneck followed by a recent and instantaneous demographic expansion that restored a large portion (>30%) of their ancestral effective population sizes shortly after introductions took place (<60 years ago). Considerable genetic admixture of some populations suggest that hypothetical barriers to dispersal (i.e., land and sea water) are permeable to gene flow and/or that they originated from introductions involving multiple lineages. This study demonstrates the repeated arrival of propagules with different origins and short time lags between arrival and establishment, emphasizing the extraordinary capacity of the species to recover from founder effects and genetically admix in invaded areas. This can explain the demonstrated capacity of this aquatic insect to spread and outcompete native species once it colonizes new suitable regions. Future genomic analyses of native range populations could help to infer the genetic makeup of introduced populations and track invasion routes.
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Substâncias Explosivas , Heterópteros , Animais , Variação Genética , Genética Populacional , Genômica , Heterópteros/genética , Espécies Introduzidas , Densidade DemográficaRESUMO
Transitions from fresh to saline habitats are restricted to a handful of insect lineages, as the colonization of saline waters requires specialized mechanisms to deal with osmotic stress. Previous studies have suggested that tolerance to salinity and desiccation could be mechanistically and evolutionarily linked, but the temporal sequence of these adaptations is not well established for individual lineages. We combined molecular, physiological and ecological data to explore the evolution of desiccation resistance, hyporegulation ability (i.e., the ability to osmoregulate in hyperosmotic media) and habitat transitions in the water beetle genus Enochrus subgenus Lumetus (Hydrophilidae). We tested whether enhanced desiccation resistance evolved before increases in hyporegulation ability or vice versa, or whether the two mechanisms evolved in parallel. The most recent ancestor of Lumetus was inferred to have high desiccation resistance and moderate hyporegulation ability. There were repeated shifts between habitats with differing levels of salinity in the radiation of the group, those to the most saline habitats generally occurring more rapidly than those to less saline ones. Significant and accelerated changes in hyporegulation ability evolved in parallel with smaller and more progressive increases in desiccation resistance across the phylogeny, associated with the colonization of meso- and hypersaline waters during global aridification events. All species with high hyporegulation ability were also desiccation-resistant, but not vice versa. Overall, results are consistent with the hypothesis that desiccation resistance mechanisms evolved first and provided the physiological basis for the development of hyporegulation ability, allowing these insects to colonize and diversify across meso- and hypersaline habitats.
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Aclimatação , Evolução Biológica , Besouros/genética , Besouros/fisiologia , Desidratação , Tolerância ao Sal , Animais , Ecossistema , Osmorregulação , FilogeniaRESUMO
Exposing organisms to a particular stressor may enhance tolerance to a subsequent stress, when protective mechanisms against the two stressors are shared. Such cross-tolerance is a common adaptive response in dynamic multivariate environments and often indicates potential co-evolution of stress traits. Many aquatic insects in inland saline waters from Mediterranean-climate regions are sequentially challenged with salinity and desiccation stress. Thus, cross-tolerance to these physiologically similar stressors could have been positively selected in insects of these regions. We used adults of the saline water beetles Enochrus jesusarribasi (Hydrophilidae) and Nebrioporus baeticus (Dytiscidae) to test cross-tolerance responses to desiccation and salinity. In independent laboratory experiments, we evaluated the effects of (i) salinity stress on the subsequent resistance to desiccation and (ii) desiccation stress (rapid and slow dehydration) on the subsequent tolerance to salinity. Survival, water loss and haemolymph osmolality were measured. Exposure to stressful salinity improved water control under subsequent desiccation stress in both species, with a clear cross-tolerance (enhanced performance) in N. baeticus In contrast, general negative effects on performance were found under the inverse stress sequence. The rapid and slow dehydration produced different water loss and haemolymph osmolality dynamics that were reflected in different survival patterns. Our finding of cross-tolerance to salinity and desiccation in ecologically similar species from distant lineages, together with parallel responses between salinity and thermal stress previously found in several aquatic taxa, highlights the central role of adaption to salinity and co-occurring stressors in arid inland waters, having important implications for the species' persistence under climate change.
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Besouros/fisiologia , Dessecação , Tolerância ao Sal , Adaptação Fisiológica , Animais , Hemolinfa/fisiologia , Pressão Osmótica , Salinidade , Estresse FisiológicoRESUMO
Geographical ranges vary greatly in size and position, even within recent clades, but the factors driving this remain poorly understood. In aquatic beetles, thermal niche has been shown to be related to both the relative range size and position of congeners but whether other physiological parameters play a role is unknown. Metabolic plasticity may be critical for species occupying more variable thermal environments and maintaining this plasticity may trade-off against other physiological processes such as immunocompetence. Here we combine data on thermal physiology with measures of metabolic plasticity and immunocompetence to explore these relationships in Deronectes (Dytiscidae). While variation in latitudinal range extent and position was explained in part by thermal physiology, aspects of metabolic plasticity and immunocompetence also appeared important. Northerly distributed, wide-ranging species apparently used different energy reserves under thermal stress from southern endemic congeners and differed in their antibacterial defences. This is the first indication that these processes may be related to geographical range, and suggests parameters that may be worthy of exploration in other taxa.
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Besouros/metabolismo , Distribuição Animal , Animais , Besouros/imunologia , Ecossistema , Europa (Continente) , Geografia , Filogeografia , TemperaturaRESUMO
Biological invasions have increased significantly in response to global change and constitute one of the major causes of biodiversity loss. Insects make up a large fraction of invasive species, in general, and freshwaters are among the most invaded ecosystems on our planet. However, even though aquatic insects dominate most inland waters, have unparalleled taxonomic diversity and occupy nearly all trophic niches, there are almost no invasive insects in freshwaters. We present some hypotheses regarding why aquatic insects are not common among aquatic invasive organisms, suggesting that it may be the result of a suite of biological, ecological and anthropogenic factors. Such specific knowledge introduces a paradox in the current scientific discussion on invasive species; therefore, a more in-depth understanding could be an invaluable aid to disentangling how and why biological invasions occur.
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Água Doce , Insetos/fisiologia , Espécies Introduzidas , Animais , Organismos Aquáticos , EcossistemaRESUMO
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.
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Ecossistema , Atividades Humanas , Insetos/fisiologia , Modelos Biológicos , Rios/química , Agricultura , Animais , Aquecimento Global , Humanos , Salinidade , Estresse Fisiológico , Água/químicaRESUMO
Salinity is one of the most important drivers of the distribution, abundance and diversity of organisms. Previous studies on the evolution of saline tolerance have been mainly centred on marine and terrestrial organisms, while lineages inhabiting inland waters remain largely unexplored. This is despite the fact that these systems include a much broader range of salinities, going from freshwater to more than six times the salinity of the sea (i.e. >200 g/L). Here, we study the pattern and timing of the evolution of the tolerance to salinity in an inland aquatic lineage of water beetles (Enochrus species of the subgenus Lumetus, family Hydrophilidae), with the general aim of understanding the mechanisms by which it was achieved. Using a time-calibrated phylogeny built from five mitochondrial and two nuclear genes and information about the salinity tolerance and geographical distribution of the species, we found that salinity tolerance appeared multiple times associated with periods of global aridification. We found evidence of some accelerated transitions from freshwater directly to high salinities, as reconstructed with extant lineages. This, together with the strong positive correlation found between salinity tolerance and aridity of the habitats in which species are found, suggests that tolerance to salinity may be based on a co-opted mechanism developed originally for drought resistance.
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Evolução Biológica , Besouros/genética , Tolerância ao Sal/genética , Animais , Núcleo Celular/genética , Besouros/fisiologia , DNA Mitocondrial/genética , Ecossistema , Modelos Genéticos , Filogenia , Salinidade , Análise de Sequência de DNARESUMO
Flight dispersal is ecologically relevant for the survival of supratidal rockpool insects. Dispersal has important consequences for colonisation, gene flow, and evolutionary divergence. Here, we compared the flight dispersal capacity of two congeneric beetle species (Ochthebius quadricollis and Ochthebius lejolisii) that exclusively inhabit these temporary, fragmented, and extreme habitats. We estimated flight capacity and inferred dispersal in both species using different approaches: experimental flying assays, examination of wing morphology, and comparison of microsatellite markers between species. Our findings revealed that both species exhibited similar flight behaviour, with 60 to 80% of the individuals flying under water heating conditions. Notably, females of both species had larger body sizes and wing areas, along with lower wing loading, than males in O. quadricollis. These morphological traits are related to higher dispersal capacity and more energetically efficient flight, which could indicate a female-biassed dispersal pattern. The wing shapes of both species are characterised by relatively larger and narrower wings in relation to other species of the genus, suggesting high flight capacity at short distances. Molecular data revealed in both species low genetic divergences between neighbouring populations, non-significant differences between species, and no isolation by distance effect at the study scale (<100 km). These results point to passive dispersal assisted by wind.
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For most nanopositioning systems, maximizing positioning bandwidth to accurately track periodic and aperiodic reference signals is the primary performance goal. Closed-loop control schemes are employed to overcome the inherent performance limitations such as mechanical resonance, hysteresis and creep. Most reported control schemes are integer-order and combine both damping and tracking actions. In this work, fractional-order controllers from the positive position feedback family namely: the Fractional-Order Integral Resonant Control (FOIRC), the Fractional-Order Positive Position Feedback (FOPPF) controller, the Fractional-Order Positive Velocity and Position Feedback (FOPVPF) controller and the Fractional-Order Positive, Acceleration, Velocity and Position Feedback (FOPAVPF) controller are designed and analysed. Compared with their classical integer-order implementation, the fractional-order damping and tracking controllers furnish additional design (tuning) parameters, facilitating superior closed-loop bandwidth and tracking accuracy. Detailed simulated experiments are performed on recorded frequency-response data to validate the efficacy, stability and robustness of the proposed control schemes. The results show that the fractional-order versions deliver the best overall performance.
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Stictonectes abellani sp. n. is described from the Iberian Peninsula. On average, the new species is larger and the colouration of the upper surface darker than in most other species of the genus. Seemingly the species has been confounded with others in the past, particularly S. optatus (Seidlitz, 1887). Males can be separated from externally similar species by studying the shape of the parameres. Additionally, the anterior margin of the clypeus is provided with a distinct rim in both sexes, which is absent or only weakly present in other species. The habitus and the male genitalia of the new species are illustrated, and compared with those of S. optatus. External morphological differences from other members of the genus are discussed. According to studies of the molecular phylogeny, based on fragments of four mitochondrial genes, S. abellani sp. n. is clearly separated from previously described species of Stictonectes Brinck, 1943, apparently being relatively basal within the genus. The new species is rather widely distributed in the south-western part of the Iberian Peninsula, inhabiting pools in small temporary siliceous streams. We provide distributional maps for all eight Iberian Stictonectes and estimate the potential distributional areas of the new species and the other two endemic Iberian species S. occidentalis Fresneda & Fery, 1990 and S. rebeccae Bilton, 2011, based on environmental niche modelling.
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Distribuição Animal , Besouros/classificação , Ecossistema , Estruturas Animais/anatomia & histologia , Animais , Besouros/anatomia & histologia , Besouros/genética , Europa (Continente) , Feminino , Masculino , Dados de Sequência Molecular , FilogeniaRESUMO
Here we focus on designing, for the first time, microsatellite markers for evolutionary and ecological research on aquatic beetles from the genus Ochthebius (Coleoptera, Hydraenidae). Some of these non-model species, with high cryptic diversity, exclusively inhabit supratidal rockpools, extreme and highly dynamic habitats with important anthropogenic threats. We analysed 15 individuals of four species (O. lejolisii, O. subinteger, O. celatus, and O. quadricollis) across 10 localities from the Mediterranean coasts of Spain and Malta. Using next-generation sequencing technology, two libraries were constructed to interpret the species of the two subgenera present consistently (Ochthebius s. str., O. quadricollis; and Cobalius, the rest of the species). Finally, 20 markers (10 for each subgenus) were obtained and successfully tested by cross-validation in the four species under study. As a by-catch, we could retrieve the complete mitochondrial genomes of O. lejolisii, O. quadricollis, and O. subinteger. Interestingly, the mitochondrial genome of O. quadricollis exhibited high genetic variability compared to already published data. The novel SSR panels and mitochondrial genomes for Ochthebius will be valuable in future research on species identification, diversity, genetic structure, and population connectivity in highly dynamic and threatened habitats such as supratidal coastal rockpools.
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In the context of aridification in Mediterranean regions, desiccation resistance and physiological plasticity will be key traits for the persistence of aquatic insects exposed to increasing desiccation stress. Control of cuticular transpiration through changes in the quantity and composition of epicuticular hydrocarbons (CHCs) is one of the main mechanisms of desiccation resistance in insects, but it remains largely unexplored in aquatic ones. We studied acclimation responses to desiccation in adults of two endemic water beetles from distant lineages living in Mediterranean intermittent saline streams: Enochrus jesusarribasi (Hydrophilidae) and Nebrioporus baeticus (Dytiscidae). Cuticular water loss and CHC composition were measured in specimens exposed to a prior non-lethal desiccation stress, allowed to recover and exposed to a subsequent desiccation treatment. E. jesusarribasi showed a beneficial acclimation response to desiccation: pre-desiccated individuals reduced cuticular water loss rate in a subsequent exposure by increasing the relative abundance of cuticular methyl-branched compounds, longer chain alkanes and branched alkanes. In contrast, N. baeticus lacked acclimation capacity for controlling water loss and therefore may have a lower physiological capacity to cope with increasing aridity. These results are relevant to understanding biochemical adaptations to drought stress in inland waters in an evolutionary and ecological context.
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Dispersal is an essential process in population and community dynamics, but is difficult to measure in the field. In freshwater ecosystems, information on biological traits related to organisms' morphology, life history and behaviour provides useful dispersal proxies, but information remains scattered or unpublished for many taxa. We compiled information on multiple dispersal-related biological traits of European aquatic macroinvertebrates in a unique resource, the DISPERSE database. DISPERSE includes nine dispersal-related traits subdivided into 39 trait categories for 480 taxa, including Annelida, Mollusca, Platyhelminthes, and Arthropoda such as Crustacea and Insecta, generally at the genus level. Information within DISPERSE can be used to address fundamental research questions in metapopulation ecology, metacommunity ecology, macroecology and evolutionary ecology. Information on dispersal proxies can be applied to improve predictions of ecological responses to global change, and to inform improvements to biomonitoring, conservation and management strategies. The diverse sources used in DISPERSE complement existing trait databases by providing new information on dispersal traits, most of which would not otherwise be accessible to the scientific community.
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Distribuição Animal , Organismos Aquáticos , Invertebrados , Animais , Conservação dos Recursos Naturais , Ecologia , Monitoramento Ambiental , Europa (Continente)RESUMO
Salinity tolerance has enabled the colonization of inland saline waters and promoted species diversification in some lineages of aquatic insects. However, the mechanisms behind this tolerance, particularly the role of cuticle hydrocarbons (CHCs), are not well-known. We characterized the CHC profile of eight species of two water beetle genera (Nebrioporus, Adephaga: Dytiscidae and Enochrus, Polyphaga: Hydrophilidae), which span the fresh-hypersaline gradient, to: i) determine the interspecific variation of CHC composition in relation to species' salinity tolerance; ii) explore plastic adjustments in CHC profiles in response to salinity changes at the intraspecific level in saline-tolerant species. CHC profiles were highly species-specific, more complex and diverse in composition, and characterized by longer-chain-length compounds in the species with higher salinity tolerance within each genus. Higher salinity tolerance in the Enochrus species was also associated with an increase in the relative abundance of branched alkanes, and with a lower proportion of n-alkanes and unsaturated compounds. These CHC characteristics are related with improved waterproofing capacity and suggest that reducing cuticle permeability was one of the key mechanisms to adapt to saline waters. Similar CHC composition patterns were found at the intraspecific level between populations from lower and higher salinity sites within saline-tolerant species of each genus. These saline species also displayed an extraordinary ability to adjust CHC profiles to changing salinity conditions in the laboratory in a relatively short time, which reflects great plasticity and a high potential to deal with daily and seasonal environmental fluctuations in the highly dynamic saline habitats.
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Besouros/metabolismo , Hidrocarbonetos/metabolismo , Tolerância ao Sal , Água/fisiologia , Aclimatação , Animais , Pressão Osmótica , Especificidade da EspécieRESUMO
Naturally stressed ecosystems hold a unique fraction of biodiversity. However, they have been largely ignored in biomonitoring and conservation programmes, such as the EU Water Framework Directive, while global change pressures are threatening their singular values. Here we present a framework to classify and evaluate the ecological quality of naturally stressed rivers along a water salinity gradient. We gathered datasets, including aquatic macroinvertebrate assemblages and environmental information, for 243 river locations across the western Mediterranean to: a) gauge the role of natural stressors (salinity) in driving aquatic community richness and composition; b) make river classifications by encompassing the wide range of environmental and biological variation exhibited by Mediterranean rivers; c) provide effective biomonitoring metrics of ecological quality for saline rivers. Our results showed that water salinity played a pivotal role in explaining the community richness and compositional changes in rivers, even when considering other key and commonly used descriptors, such as elevation, climate or lithology. Both environmental and biologically-based classifications included seven river types: three types of freshwater perennial rivers, one freshwater intermittent river type and three new saline river types. These new saline types were not included in previous classifications. Their validation by independent datasets showed that the saline and freshwater river types represented differentiable macroinvertebrate assemblages at family and species levels. Biomonitoring metrics based on the abundance of indicator taxa of each saline river type provided a much better assessment of the ecological quality of saline rivers than other widely used biological metrics and indices. Here we demonstrate that considering natural stressors, such as water salinity, is essential to design effective and accurate biomonitoring programmes for rivers and to preserve their unique biodiversity.
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Biodiversidade , Ecossistema , Monitoramento Ambiental/métodos , Rios/química , Salinidade , Animais , Itália , Marrocos , EspanhaRESUMO
The fast and accurate tracking of periodic and arbitrary reference trajectories is the principal goal in many nanopositioning applications. Flexure-based piezoelectric stack driven nanopositioners are widely employed in applications where accurate mechanical displacements at these nanometer scales are required. The performance of these nanopositioners is limited by the presence of lightly damped resonances in their dynamic response and actuator nonlinearities. Closed-loop control techniques incorporating both damping and tracking are typically used to address these limitations. However, most tracking schemes employed use a first-order integrator where a triangular trajectory commonly used in nanopositioning applications necessitates a double integral for zero-error tracking. The phase margin of the damped system combined with the hardware-induced delay deem the implementation of a double-integrator unstable. To overcome this limitation, this paper presents the design, analysis and application of a new control scheme based on the structure of the traditional Two-Degrees-of-Freedom PID controller (2DOF-PID). The proposed controller replaces the integral action of the traditional 2DOF-PID with a double integral action (2DOF-PI2D). Despite its simplicity, the proposed controller delivers superior tracking performance compared to traditional combined damping and tracking control schemes based on well-reported designs such as positive position feedback (PPF), Integral resonant control (IRC), and Positive Velocity and Position Feedback (PVPF). The stability of the control system is analyzed in the presence of a time delay in the system. Experimental results validating the efficacy of the proposed chattering-free control of a piezo-driven nanopositioning system are included.
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New faunistic and distributional data on aquatic Hemiptera from the Oriental Region of Morocco and the Moulouya River basin, covering an area of 119,268 km2, are presented. An annotated list of 45 species, 21 genera and 11 families is provided. Nine species are new records for the entire studied area, and two for the Moulouya River basin. We rediscovered Velia noualhieri iberica and Notonecta viridis after eight decades. We also provide novel records for Aphelocheirus pemae and Hebrus atlas, which were recently described from Morocco, and for the invasive corixid Trichocorixa verticalis verticalis. The dominance of species with broad distributions points to the high dispersal dynamics of the Hemiptera community in the study area, similar to that of other areas where a semiarid climate prevails.
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Heterópteros , Animais , Clima , Marrocos , RiosRESUMO
By exploiting the co-located sensor-actuator arrangement in typical flexure-based piezoelectric stack actuated nanopositioners, the polezero interlacing exhibited by their axial frequency response can be transformed to a zero-pole interlacing by adding a constant feed-through term. The Integral Resonant Control (IRC) utilizes this unique property to add substantial damping to the dominant resonant mode by the use of a simple integrator implemented in closed loop. IRC used in conjunction with an integral tracking scheme, effectively reduces positioning errors introduced by modelling inaccuracies or parameter uncertainties. Over the past few years, successful application of the IRC control technique to nanopositioning systems has demonstrated performance robustness, easy tunability and versatility. The main drawback has been the relatively small positioning bandwidth achievable. This paper proposes a fractional order implementation of the classical integral tracking scheme employed in tandem with the IRC scheme to deliver damping and tracking. The fractional order integrator introduces an additional design parameter which allows desired pole-placement, resulting in superior closed loop bandwidth. Simulations and experimental results are presented to validate the theory. A 250% improvement in the achievable positioning bandwidth is observed with proposed fractional order scheme.