{"title":"Effective dispersal of fern spore and the ecological relevance of zoochory.","authors":"James M R Brock","doi":"10.1111/brv.70038","DOIUrl":"https://doi.org/10.1111/brv.70038","url":null,"abstract":"<p><p>The mechanisms of fern dispersal are under-studied and there are few data to support the vectors assumed to drive patterns of sporophyte occurrence and speciation. Although wind is generally the fern spore dispersal vector described in the literature, there has always been competing evidence supporting alternate vectors. Despite this, established patterns of dispersal are generally discussed in the context of wind. The assumptions around wind as a dispersal vector and the possibilities of single-spore establishment have confounded discussions on effective dispersal of fern spore, fern meta-population dynamics, and fern speciation. In this study, I review (i) the importance of spore load across taxa, (ii) evidence for vectors of fern spore, (iii) the environmental tolerances of fern life stages, and (iv) the relevance of, and constraints on, different dispersal vectors in the context of increasingly hazardous landscape matrices. I conclude that whilst wind is an important dispersal vector in non-hazardous landscapes, directed dispersal by an animal vector to isolated safe sites in a hazardous landscape matrix may be key for fern metapopulations and communities.</p>","PeriodicalId":133,"journal":{"name":"Biological Reviews","volume":" ","pages":""},"PeriodicalIF":11.0,"publicationDate":"2025-05-29","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"144172243","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":1,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Irene Sánchez González, Garrett W Hopper, Jamie R Bucholz, Jeffrey D Lozier, Carla L Atkinson
{"title":"A functional morphological classification for North American freshwater mussels: associations between morphology and environmental parameters across spatial scales.","authors":"Irene Sánchez González, Garrett W Hopper, Jamie R Bucholz, Jeffrey D Lozier, Carla L Atkinson","doi":"10.1111/brv.70040","DOIUrl":"https://doi.org/10.1111/brv.70040","url":null,"abstract":"<p><p>Trait-based approaches have emerged as a general framework that translates species-specific knowledge to understand the processes driving patterns of diversity and distributions. Morphological traits are relatively easy to measure and can provide information on organism-environment interactions and community structure due to their close linkage to ecological function and habitat partitioning. Freshwater mussels (Family: Unionidae) are a diverse (~360 North American species) and endangered group of organisms. Mussels display great interspecific morphological variation potentially yielding broad ecological implications. We aimed to modify quantitively an existing shell morphological classification system by combining size, shape, and sculpturing data using a robust data set of 1362 individuals representing 64 species spanning a broad cross section of the diverse North American freshwater mussel fauna. Using multivariate techniques, we classified species into morphological classes based on trait similarities hypothesized to explain species distributions and habitat associations. We then tested how well the classification system predicted trait-environment relationships using quantitative mussel survey data with paired environmental data collected at three spatial scales [river (km), reach (40-150 m), patch (0.25 m<sup>2</sup>)]. Mussel species clustered into six different morphological classes based on sculpturing, shape, and body size traits. We found associations between morphological classes and environmental parameters at each spatial scale. The modified classification explained more variation in community distribution as predicted by abiotic variables than previous frameworks. Our study underscores the value of morphological traits in predicting species distributions and understanding mechanisms of community assembly and we provide a foundation for fellow researchers to expand our morphological classification. This knowledge has significant implications for mussel conservation and management, as it helps identify suitable habitats that can guide reintroduction strategies through incorporating multiple spatial scales, a broad representation of species and geographical distribution and a wide suite of morphological traits.</p>","PeriodicalId":133,"journal":{"name":"Biological Reviews","volume":" ","pages":""},"PeriodicalIF":11.0,"publicationDate":"2025-05-29","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"144172241","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":1,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"The biogeographic and genomic signatures of dynamic river networks for terrestrial species in Amazonia.","authors":"Lukas J Musher","doi":"10.1111/brv.70042","DOIUrl":"https://doi.org/10.1111/brv.70042","url":null,"abstract":"<p><p>Amazonia contains Earth's largest freshwater basin, largest contiguous stretch of tropical forest, and most species-rich terrestrial biota on Earth. Rivers are key geographic features that drive diversification of the Amazonian biota, but they are also dynamic, which challenges their role as long-term barriers to dispersal and gene flow. The impacts of such river dynamics on organismal evolution have only recently been explored in detail. Here I examine biodiversity patterns and processes in Amazonia to elucidate how taxa diversify in the context of river network dynamics. I borrow the River Capture Hypothesis from ichthyology, and draw on evidence from speciation genomics, hybrid zones, and community assembly to demonstrate the effects of river network evolution on biodiversification. The idea is simple: populations of organisms whose dispersal is restricted by rivers become semi-isolated by rivers. Drift and selection against introgression drive divergence, but as rivers move, previously isolated populations come into secondary contact, facilitating lineage fusions or the migration of hybrid zones to other rivers. The basin's unique macroecological patterns and rich biota thus may have resulted from repeated divergences, lineage fusions, and range expansions around a network of non-stationary extrinsic barriers with variable results depending on the degree of intrinsic reproductive isolation that accumulates during this process. The evolutionary consequences of dynamic landscapes extend beyond Amazonia as \"fission-fusion-fission\" cycles modulate the diversification and spatial patterning of life on Earth in general.</p>","PeriodicalId":133,"journal":{"name":"Biological Reviews","volume":" ","pages":""},"PeriodicalIF":11.0,"publicationDate":"2025-05-26","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"144148891","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":1,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Synergy, complexity, and the dirty, dirty cheats of the world.","authors":"Jennifer A H Koop, Neil W Blackstone","doi":"10.1111/brv.70041","DOIUrl":"https://doi.org/10.1111/brv.70041","url":null,"abstract":"<p><p>Studies of symbiosis employ the term \"parasitism\" to connote different sorts of relationships. Within the context of mutualistic symbioses, parasites are otherwise cooperative individuals or strains that appropriate a disproportionate amount of the synergistic products. In the context of antagonistic symbioses, there is no pretence of cooperation, and instead parasites are defined as individuals or strains that derive fitness benefits at a fitness cost to their hosts. In both cases, parasitism is selected for at the lower level (that of the individual symbiont) but selected against at the higher level (the group of symbionts in a single host). Despite these similarities, these different sorts of parasitism likely evolve by different pathways. Once a host-symbiont relationship initiates, if functional synergy is lacking, the relationship will remain exploitative, although parasites may differ in their detrimental effects on the host and the higher-level unit. If functional synergy is present, however, cooperation may develop with benefits for both host and symbionts (i.e. mutualism). Nevertheless, parasites may still evolve from within these incipient relationships when individuals or strains of symbionts act parasitically by defecting from the common good to further their selfish replication. Levels-of-selection dynamics thus underlie both forms of parasitism, but only in the case of latent functional synergy can true symbiotic complexity at the higher level emerge.</p>","PeriodicalId":133,"journal":{"name":"Biological Reviews","volume":" ","pages":""},"PeriodicalIF":11.0,"publicationDate":"2025-05-26","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"144141055","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":1,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Infection dynamics of endosymbionts that manipulate arthropod reproduction.","authors":"Franziska A Brenninger, Roman Zug, Hanna Kokko","doi":"10.1111/brv.70024","DOIUrl":"https://doi.org/10.1111/brv.70024","url":null,"abstract":"<p><p>A large proportion of arthropod species are infected with endosymbionts, some of which selfishly alter host reproduction. The currently known forms of parasitic reproductive manipulations are male-killing, feminization, cytoplasmic incompatibility, parthenogenesis induction and distortion of sex allocation. While all of these phenomena represent adaptations that enhance parasite spread, they differ in the mechanisms involved and the consequent infection dynamics. We focus here on the latter aspect, summarizing existing theoretical literature on infection dynamics of all known reproductive manipulation types, and completing the remaining knowledge gaps where dynamics have not been modelled yet. Our unified framework includes the minimal model components required to describe the effects of each manipulation. We establish invasion criteria for all potential combinations of manipulative endosymbionts, yielding predictions for an endosymbiont's increase from rarity within a host population that is initially either uninfected or infected with a different symbiont strain. We consider diplodiploid and haplodiploid hosts, as the mechanisms as well as the infection dynamics of reproductive manipulations can differ between them. Our framework reveals that endosymbionts that a priori have the best invasion prospects are not necessarily the most commonly found ones in nature; priority effects play a role too, and cytoplasmic incompatibility excels in this regard. As a whole, considerations of the ease with which a symbiont spreads have to be complemented with knowledge of how easy it is to achieve a particular manipulation, and with factors influencing the probability that interspecific host switching occurs and succeeds.</p>","PeriodicalId":133,"journal":{"name":"Biological Reviews","volume":" ","pages":""},"PeriodicalIF":11.0,"publicationDate":"2025-05-22","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"144118396","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":1,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Marjolein E M Toorians, T Jonathan Davies, Ailene MacPherson
{"title":"Multi-host pathogen transmission and the disease-diversity relationship.","authors":"Marjolein E M Toorians, T Jonathan Davies, Ailene MacPherson","doi":"10.1111/brv.70027","DOIUrl":"https://doi.org/10.1111/brv.70027","url":null,"abstract":"<p><p>How diseases are transmitted within a multi-host community is a complex biological process with important ecological and societal consequences. The intricacies of interspecific disease transmission determine when a disease can spread to a novel host, including humans (zoonosis), and the severity of emerging epidemics. Interspecific disease transmission also mediates long-term disease prevalence within a multi-host community which is at the core of the disease-diversity relationship. Mathematical models play a central role in formulating predictions about spillover, prevalence, and the disease-diversity relationship. Yet, how the complexity of transmission is captured (or not) by the assumptions of these models is often unclear. Here, we decompose the transmission process into five biological stages using bovine tuberculosis (bTB) as an illustrative example of transmission in a multi-host system. We then examine the often-implicit assumptions that classic compartmental models make about this process. We use the intuition gained from this decomposition to formulate hypotheses for how transmission can mediate outbreak potential, infection prevalence, and the amplifying or diluting effects of host diversity on disease prevalence. We further illustrate the key principles and implications of transmission with a diverse array of examples of multi-host pathogens. Throughout we emphasise the role of evolution in shaping interspecific transmission, from the evolutionary relatedness of the hosts themselves to the adaptation of the pathogen to novel hosts.</p>","PeriodicalId":133,"journal":{"name":"Biological Reviews","volume":" ","pages":""},"PeriodicalIF":11.0,"publicationDate":"2025-05-15","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"144075057","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":1,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"The pace and shape of ant ageing.","authors":"Luisa M Jaimes-Nino, Jan Oettler","doi":"10.1111/brv.70035","DOIUrl":"https://doi.org/10.1111/brv.70035","url":null,"abstract":"<p><p>Ants have been proposed as good models to study ageing and the effects of extrinsic mortality because of their long lifespans and plasticity of ageing within species. We discuss how age-dependent extrinsic mortality might influence queen lifespan, and how the effect of age-independent extrinsic mortality needs further study, accounting for different density-dependence scenarios. Based on a critical review of the available demographic data, we discuss the selective forces underlying ant ageing. We discuss differences and similarities between the life-history strategy of ants and the reproductive strategies iteroparity and semelparity. We consider how late-life fitness gains for the \"superorganism\" select for a delay of actuarial, and reproductive senescence, and we suggest future research directions.</p>","PeriodicalId":133,"journal":{"name":"Biological Reviews","volume":" ","pages":""},"PeriodicalIF":11.0,"publicationDate":"2025-05-15","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"144075062","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":1,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"The evolution of reproduction in Ediacaran-Cambrian metazoans.","authors":"Rachel A Wood, Mary L Droser","doi":"10.1111/brv.70036","DOIUrl":"https://doi.org/10.1111/brv.70036","url":null,"abstract":"<p><p>The evolution of reproductive style is a fundamental aspect of metazoan life history but has not been explored holistically through the Ediacaran-Cambrian rise of metazoans. Recent molecular clock analyses based on only unequivocal metazoan fossil calibrations suggest that Porifera were present by at least 590 million years ago (Ma), all major eumetazoan clades originated in the mid-late Ediacaran, and bilaterians were probably present by the late Ediacaran. An alternating pelagic larval (potentially for dispersal) and benthic adult life cycle appears to be an ancestral feature of metazoans. A compilation of inferred reproductive styles from the fossil record reveals that the low-competition, deep-water communities of the Ediacaran Avalon macrofossil assemblage (ca. 575 to 560 Ma) had current-borne sexually produced larval with both local (non-planktotrophic, with no feeding) and more widespread (planktotrophic, with feeding) dispersal followed by vegetative growth. By ca. 560 Ma, White Sea assemblage communities in shallow settings show dense aggregations, which were often dominated by single populations of episodic sexually produced larval spatfalls. Some taxa may show potential larval philopatry. By 550 Ma, with the rise of biomineralization and colonisation of shallow marine carbonate settings, the ability to encrust hard substrates, create multiple branches via budding, and rudimentary mutual attachment of inferred clones, first appear. The dominant apparent mode of reproduction throughout the Ediacaran was therefore via current-borne sexually produced larvae followed by asexual reproduction, via either budding, fragmentation or fission. In these communities where biotic interactions were limited, this enabled colonisation of newly available soft and hard substrates followed by rapid growth. Early Cambrian communities showed increased endemism, enhanced trophic interactions and widespread macropredation. By the early Cambrian Fortunian stage (ca. 535 Ma), gonochorism (separate sexes) may have been present in priapulid worms. During Cambrian Stage 2 (ca. 532 Ma), internal fertilisation probably appeared in molluscs but widespread planktotrophy did not appear until the latest Cambrian/early Ordovician. Mutual attachment of diverse skeletal taxa became more common, particularly within reefs. Evidence for egg brooding and parental care in arthropods had appeared by the early Stage 3 (ca. 518 Ma). While reproductive styles were independently acquired, this overall pattern suggests a shift both to higher fecundity and to higher quality offspring in some groups during the Ediacaran-Cambrian Radiation, driven by increasing biotic interactions, including the rise of macropredation.</p>","PeriodicalId":133,"journal":{"name":"Biological Reviews","volume":" ","pages":""},"PeriodicalIF":11.0,"publicationDate":"2025-05-15","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"144075059","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":1,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Juan Olvido Perea-García, Aurora Teuben, Kai R Caspar
{"title":"Look past the cooperative eye hypothesis: reconsidering the evolution of human eye appearance.","authors":"Juan Olvido Perea-García, Aurora Teuben, Kai R Caspar","doi":"10.1111/brv.70033","DOIUrl":"https://doi.org/10.1111/brv.70033","url":null,"abstract":"<p><p>The external appearance of the human eye has been prominently linked to the evolution of complex sociocognitive functions in our species. The cooperative eye hypothesis (CEH) proposes that human eyeballs, with their weakly expressed conjunctival and scleral pigmentation, are uniquely conspicuous and evolved under selective pressures to behave cooperatively, therefore signalling attentiveness to conspecifics. Non-human primates are instead assumed to display less-salient eye morphologies that help mask their gaze to facilitate competitive, rather than cooperative actions. Here, we argue that the CEH, although continuing to be influential, lacks robust empirical support. Over the past two decades, multidisciplinary research has undermined its original rationale and central premises: human eye pigmentation does not uniquely stand out among primates, it is not uniform at species level and the available evidence does not conclusively suggest that it facilitates gaze following to notable extents. Hence, the CEH currently provides a theoretical framework that risks confusing, rather than informing, inferences about the evolution of human external eye appearance and its selective drivers. In a call to move past it, we review alternative hypotheses with the potential to elucidate the emergence of the human ocular phenotype from the considerable spectrum of diversity found within the primate order.</p>","PeriodicalId":133,"journal":{"name":"Biological Reviews","volume":" ","pages":""},"PeriodicalIF":11.0,"publicationDate":"2025-05-14","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"143959061","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":1,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Nadine Praeg, Michael Steinwandter, Davnah Urbach, Mark A Snethlage, Rodrigo P Alves, Martha E Apple, Peter Bilovitz, Andrea J Britton, Estelle P Bruni, Ting-Wen Chen, Kenneth Dumack, Fernando Fernandez-Mendoza, Michele Freppaz, Beat Frey, Nathalie Fromin, Stefan Geisen, Martin Grube, Elia Guariento, Antoine Guisan, Qiao-Qiao Ji, Juan J Jiménez, Stefanie Maier, Lucie A Malard, Maria A Minor, Cowan C Mc Lean, Edward A D Mitchell, Thomas Peham, Roberto Pizzolotto, Andy F S Taylor, Philippe Vernon, Johan J van Tol, Donghui Wu, Yunga Wu, Zhijing Xie, Bettina Weber, Paul Illmer, Julia Seeber
{"title":"Biodiversity in mountain soils above the treeline.","authors":"Nadine Praeg, Michael Steinwandter, Davnah Urbach, Mark A Snethlage, Rodrigo P Alves, Martha E Apple, Peter Bilovitz, Andrea J Britton, Estelle P Bruni, Ting-Wen Chen, Kenneth Dumack, Fernando Fernandez-Mendoza, Michele Freppaz, Beat Frey, Nathalie Fromin, Stefan Geisen, Martin Grube, Elia Guariento, Antoine Guisan, Qiao-Qiao Ji, Juan J Jiménez, Stefanie Maier, Lucie A Malard, Maria A Minor, Cowan C Mc Lean, Edward A D Mitchell, Thomas Peham, Roberto Pizzolotto, Andy F S Taylor, Philippe Vernon, Johan J van Tol, Donghui Wu, Yunga Wu, Zhijing Xie, Bettina Weber, Paul Illmer, Julia Seeber","doi":"10.1111/brv.70028","DOIUrl":"https://doi.org/10.1111/brv.70028","url":null,"abstract":"<p><p>Biological diversity in mountain ecosystems has been increasingly studied over the last decade. This is also the case for mountain soils, but no study to date has provided an overall synthesis of the current state of knowledge. Here we fill this gap with a first global analysis of published research on cryptogams, microorganisms, and fauna in mountain soils above the treeline, and a structured synthesis of current knowledge. Based on a corpus of almost 1400 publications and the expertise of 37 mountain soil scientists worldwide, we summarise what is known about the diversity and distribution patterns of each of these organismal groups, specifically along elevation, and provide an overview of available knowledge on the drivers explaining these patterns and their changes. In particular, we document an elevation-dependent decrease in faunal diversity above the treeline, while for cryptogams there is an initial increase above the treeline, followed by a decrease towards the nival belt. Thus, our data confirm the key role that elevation plays in shaping the biodiversity and distribution of these organisms in mountain soils. The response of prokaryote diversity to elevation, in turn, was more diverse, whereas fungal diversity appeared to be substantially influenced by plants. As far as available, we describe key characteristics, adaptations, and functions of mountain soil species, and despite a lack of ecological information about the uncultivated majority of prokaryotes, fungi, and protists, we illustrate the remarkable and unique diversity of life forms and life histories encountered in alpine mountain soils. By applying rule- as well as pattern-based literature-mining approaches and semi-quantitative analyses, we identified hotspots of mountain soil research in the European Alps and Central Asia and revealed significant gaps in taxonomic coverage, particularly among biocrusts, soil protists, and soil fauna. We further report thematic priorities for research on mountain soil biodiversity above the treeline and identify unanswered research questions. Building upon the outcomes of this synthesis, we conclude with a set of research opportunities for mountain soil biodiversity research worldwide. Soils in mountain ecosystems above the treeline fulfil critical functions and make essential contributions to life on land. Accordingly, seizing these opportunities and closing knowledge gaps appears crucial to enable science-based decision making in mountain regions and formulating laws and guidelines in support of mountain soil biodiversity conservation targets.</p>","PeriodicalId":133,"journal":{"name":"Biological Reviews","volume":" ","pages":""},"PeriodicalIF":11.0,"publicationDate":"2025-05-14","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"144075053","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":1,"RegionCategory":"生物学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}