On the generality of the latitudinal diversity gradient

The decline of biodiversity with latitude has received great attention, but both the concise pattern and the causes of the gradient are under strong debate. Most studies of the latitudinal gradient comprise only one or few organism types and are often restricted to certain region or habitat types. To test for significant variation in the gradient between organisms, habitats, or regions, a meta-analysis was conducted on nearly 600 latitudinal gradients assembled from the literature. Each gradient was characterized by two effect sizes, strength (correlation coefficient) and slope, and additionally by 14 variables describing organisms, habitats, and regions. The analysis corroborated the high generality of the latitudinal diversity decline. Gradients on regional scales were significantly stronger and steeper than on local scales, and slopes also varied with sampling grain. Both strength and slope increased with organism body mass, and strength increased with trophic level. The body mass-effect size relation varied for ecto- versus homeotherm organisms and for different dispersal types, suggesting allometric effects on energy use and dispersal ability as possible mechanisms for the body mass effect. Latitudinal gradients were weaker and less steep in freshwater than in marine or terrestrial environments and differed significantly between continents and habitat types. The gradient parameters were not affected by hemisphere or the latitudinal range covered. This analysis is the first to describe these general and significant patterns, which have important consequences for models aiming to explain the latitudinal gradient.     

Dung beetle assemblages in primary forest and disturbed habitats in a tropical dry forest landscape in western Mexico

Dung beetles (Scarabaeinae) are conspicuous components in most terrestrial ecosystems, performing important ecological functions and services. Being sensitive to several types of disturbance, they have been successfully used as indicators of habitat change. Dung beetle communities in tropical rainforests have been well studied, but considerably less information is available for tropical dry forests. In this study I sampled dung beetles in two undisturbed habitats, deciduous forest and semideciduous forest, and two disturbed habitats, secondary forest and open area habitat, in the Chamela-Cuixmala region of western Mexico. Dung beetle species with high indicator value for each habitat were identified. Beetle abundance, observed species richness and estimated species richness were similar in the three forest habitats, but significantly lower in the open area habitat. A more detailed analysis of species-specific abundances in the three forest habitats revealed some differences. Transects of one of the undisturbed habitats, the deciduous forest, were more similar to the non-adjacent transects of disturbed secondary forest, than to the adjacent undisturbed semideciduous forest transects. Unlike studies in other tropical sites that have found a decrease in equitability in Scarabaeinae assemblages between undisturbed forest and disturbed habitat (particularly open habitats), in the Chamela-Cuixmala region all four habitats showed similar low equitability in community structure, with two or three very dominant species.     

Benthic habitats do show a significant latitudinal diversity gradient: A comment on Kinlock et al. (2018)

The latitudinal diversity gradient (LDG) has been investigated for decades, with hundreds of studies focusing on different organisms, regions and habitat types. Meta‐analysis may be considered, therefore, a useful tool to explore the generality and limitations of this remarkable macroecological pattern. The first meta‐analysis exploring variations in the LDG, published by Hillebrand in 2004, revealed that the latitudinal decline in species richness seems to be indeed a general phenomenon. However, Kinlock et al. (2018, Global Ecology and Biogeography, 27, 125–141) revisited recently the challenge of synthesizing individual LDGs and indicated that the phenomenon is not ubiquitous among habitats of the marine realm. More precisely, they indicated that the phenomenon is non‐significant in the benthic habitat. Here, we suggest that the marine habitat categories used by them (i.e., benthic, coral reefs, coastal, open ocean) are not independent and that reclassifying the studies significantly alters one of their main results. By assigning the studies into benthic and pelagic categories, and additionally into coastal or oceanic zones, we show that non‐ambiguous, evolutionarily meaningful marine habitats display a significant latitudinal decline in species richness.     

Re-terrestrialization in the phylogeny of epiphytic plant lineages: Microsoroid ferns as a case study

The fern family Polypodiaceae, with over 1600 species, is not only one of the most species-rich families of ferns, but also a major contributor to the vascular epiphytic diversity throughout the tropics. Although the vast majority of species belonging to this family prefer to grow as epiphytes, several species colonize successfully rheophytic, lithophytic, and even terrestrial habitats. Here, we explore the hypothesis that non-epiphytic habitat preferences, including terrestrial growth, evolved secondarily with epiphytes being the plesiomorphic habitat preference. The results of phylogenetic analyses, based on dense taxon sampling and four chloroplast DNA regions, were integrated with divergence time estimates and ancestral character state reconstructions to test these predictions. Both fossils and secondary calibration data were incorporated to obtain divergence time estimations. The results support the prediction of multiple transitions from epiphytic/lithophytic to terrestrial/rheophytic habitats occurring mainly in the Microsoreae lineage. The change in niche preferences coincides with niche colonization opportunities created by climatic fluctuations and geographical changes during the Oligocene and Miocene periods.     

Diverse migration strategy between freshwater and seawater habitats in the freshwater eel genus Anguilla

The freshwater eels of the genus Anguilla, which are catadromous, migrate between freshwater growth habitats and offshore spawning areas. A number of recent studies, however, found examples of the temperate species Anguilla anguilla, Anguilla rostrata, Anguilla japonica, Anguilla australis and Anguilla dieffenbachii that have never migrated into fresh water, spending their entire life history in the ocean. Furthermore, those studies found an intermediate type between marine and freshwater residents, which appear to frequently move between different environments during their growth phase. The discovery of marine and brackish-water residents Anguilla spp. suggests that they do not all have to be catadromous, and it calls into question the generalized classification of diadromous fishes. There has been little available information, however, concerning migration in tropical Anguilla spp. Anguilla marmorata, shows three fluctuation patterns: (1) continuous residence in fresh water, (2) continuous residence in brackish water and (3) residence in fresh water after recruitment, while returning to brackish water. Such migratory patterns were found in other tropical species, Anguilla bicolor bicolor and Anguilla bicolor pacifica. In A. b. bicolor collected in a coastal lagoon of Indonesia, two further patterns of habitat use were found: (1) constantly living in either brackish water or sea water with no freshwater life and (2) habitat shift from fresh water to brackish water or sea water. The wide range of environmental habitat use indicates that migratory behaviour of tropical Anguilla spp. is facultative among fresh, brackish and marine waters during their growth phases after recruitment to the coastal areas. Further, the migratory behaviours of tropical Anguilla spp. appear to differ in each habitat in response to inter and intra-specific competition. The results suggest that tropical Anguilla spp. have a flexible pattern of migration, with an ability to adapt to various habitats and salinities. The ability of anguillids to reside in environments of various salinities would be a common feature between tropical and temperate species without a latitudinal cline. Thus, the migration of Anguilla spp. into fresh water is clearly not an obligatory behaviour. This evidence of geographical variability among Anguilla spp. suggests that habitat use is determined by environmental conditions in each site.     

Extinction and diversification in the Devonian Brachiopoda of New York state: No correlation with sea level?

Sea level highstand is generally considered to promote high species diversities among marine organisms through habitat expansion and global climatic amelioration, and marine regression to trigger elevated extinction rates among marine benthic organisms by habitat reduction (the species‐area effect), and among both marine and terrestrial organisms by global climatic deterioration. The Devonian is unusual in that the Late Devonian mass extinction occurs during an interval of global sea level highstand. To further explore this anomaly, the potential relationship between relative sea level and evolutionary biology is analyzed here for the Brachiopoda of the Devonian Period. Successive linear modeling reveals a total lack of correlation between relative sea level and either origination rates, extinction rates, or standing diversity among the Devonian brachiopods.     

The evolution of marine insects: phylogenetic, ecological and geographical aspects of species diversity in marine water striders

No other group of insects have been more successful in colonizing marine habitats than water striders and their allies (Heteroptera, Gerromorpha). More than 10% of the 1700 species of gerromorphan bugs are marine. Water striders have colonized the marine environment at least 14 times. The fossil records suggest that marine habitats were invaded by members of the families Veliidae and Gerridae earlier than 20-30 and 45 million years before present, respectively. Estuaries and mangrove swamps are undoubtedly the ancestral type of habitat, but water striders have diversified further in marine habitats including the surface of the open ocean (sea skaters. Halobates). Except for being obligatorily flightless, marine water striders are structurally very similar to their non-marine relatives. Physiological and behavioral rather than morphological specializations are likely to have been key innovations in the transition from limnic to marine habitats. The oldest and most species-rich clades originated in the Indo-West Pacific region. There are 3.5 times as many species of marine water striders in the Indo-West Pacific region than in the Atlantic/Caribbean/East Pacific region. This "diversity anomaly" is explained historically by region-specific differences in the origin and proliferation of clades, in paleoclimate and paleogeography, and in the propensity for dispersal between regions.     

Common and rare Swedish vertebrates — distribution and habitat preferences

Data have been compiled on the distribution, habitat preferences and population sizes of 348 vertebrates reproducing in Sweden (excluding fish) and their species richness in different habitats and regions was investigated. Furthermore, we compared the habitat preferences and distribution of rare and common vertebrates. The relative species number (corrected for area) increases from the north (the hemiarctic/boreal zone) to the south (the temperate/hemiboreal zone). The relative number of species in major habitats is highest in farmland, lakes and running waters, and possibly also in the sea. However, the absolute number of species is highest in woodland, the dominant habitat in Sweden (56% of the land area excluding sea). Within woodlands, a large proportion of species occur in southern deciduous forests, in other deciduous forests and in mixed forests, while coniferous forests are less species-rich when the habitat area is taken into account. In farmland, the most species-rich habitats are meadows and forest edges, while marshes are the most species-rich habitats amongst lakes and running waters. Nationally rare species (<1000 individuals) have a smaller European range than common species, and they also have their European distribution centres further to the south than the common species. Of the species occurring in large parts of Sweden (that is, both in the north and the south) a low proportion (4.7%) are classified as rare compared with species occurring mainly in the north (20.6% rare) or mainly in the south (31.6% rare) of Sweden. There is a positive correlation between the number of rare species and the total species number in 41 subcategory habitats, and the proportion of rare species is similar in most habitats. A more detailed analysis (including effects of both regions and habitats) suggests that the proportion of rare species in a region is a result of differences between latitudes, but also of different habitats (when corrected for latitude effects). There are differences in the proportion of rare species between the nine most species-rich orders (for example, there is a high proportion of rare species in the order Carnivora and a low proportion in the order Rodentia), but the proportion of rare species in these orders is not independent of the habitats in which they occur, making it difficult to separate the effects of habitat preferences and taxonomy on rarity. The focusing of conservation work on relatively species-rich habitats in southern and middle Sweden — such as some farmland habitats (that is, meadows and forest edges), marshes (in connection with lakes and running waters) and forests with a relatively high proportion of deciduous trees — is of high priority if the conservation of biodiversity is a main goal. These are also the habitats with a high number of rare and red-listed species.     

Terrestrial or marine species distribution model: Why not both? A case study with seabirds

Species reliant on both the terrestrial and marine realms present a challenge for conventional species distribution models (SDMs). For such species, standard single‐realm SDMs may omit key information that could result in decreased model accuracy and performance. Existing approaches to habitat suitability modeling typically do not effectively combine information from multiple realms; this methodological gap can ultimately hamper management efforts for groups such as seabirds, seals, and turtles. This study, for the first time, jointly incorporates both terrestrial information and marine information into a single species distribution model framework. We do this by sampling nearby marine conditions for a given terrestrial point and vice versa using parameters set by each species’ mean maximum foraging distance and then use standard SDM methods to generate habitat suitability predictions; therefore, our method does not rely on post hoc combination of several different models. Using three seabird species with very different ecologies, we investigate whether this new multi‐realm approach can improve our ability to identify suitable habitats for these species. Results show that incorporating terrestrial information into marine SDMs, or vice versa, generally improves model performance, sometimes drastically. However, there is considerable variability between species in the level of improvement as well as in the particular method that produces the most improvement. Our approach provides a repeatable and transparent method to combine information from multiple ecological realms in a single SDM framework. Important advantages over existing solutions include the opportunity to, firstly, easily combine terrestrial and marine information for species that forage large distances inland or out to sea and, secondly, consider interactions between terrestrial and marine variables.     

热带珊瑚礁区海参的生境选择与生态作用

珊瑚礁生态系统是一个高生产力、高生物多样性的特殊海洋生态系统,具有为生物提供栖息地、参与生物地球化学循环、防浪护岸、指示水体污染程度等生态功能。珊瑚礁生态系统的突出特点是其生境异质性很高,各种各样的生境斑块为种类繁多、习性各异的游泳和底栖生物提供栖息场所,这些礁栖生物通过参与各项生态过程而形成各种特定的功能群,共同完成重要的生态功能。在热带珊瑚礁生态系统中,海参是大型底栖动物区系的重要一员。种类繁多的海参具有各自不同的生境选择特征,通过摄食、运动等行为活动发挥着改良底质、促进有机物矿化和营养盐再生等生态作用。近几年来,全球热带海参受人类过度捕捞和珊瑚礁退化的影响而面临资源衰退、物种多样性丧失等问题,深入认识其生态学功能、加强热带海参资源保护迫在眉睫。综述了国内外热带珊瑚礁海参的基础生态学研究进展：海参对珊瑚礁生境斑块呈现显著的偏好选择特征以及种间差异和季节变动,不同生境斑块的食物质量、底质类型和水动力条件是影响海参生境偏好的重要因素;海参通过生物扰动可以改变珊瑚礁生境沉积物的含水量、渗透性、颗粒组成、再矿化率、无机营养物质释放速率以及孔隙水的化学梯度,并增加沉积物中的溶氧浓度、促进溶解...     

Evaluation of macrofaunal effects on leaf litter breakdown rates in aquatic and terrestrial habitats

Abstract Decomposition of the organic matter is a key process in the functioning of aquatic and terrestrial ecosystems, although different factors influence processing rates between and within these habitats. Most patterns were described for temperate regions, with fewer studies in tropical, warmer sites. In this study, we carried out a factorial experiment to compare processing rates of mixed species of leaf litter between terrestrial and aquatic habitats at a tropical site, using ?ne and coarse mesh cages to allow or prevent colonization by macroinvertebrates. The experiment was followed for 10 weeks, and loss of leaf litter mass through time was evaluated using exponential models. We found no interaction between habitat and mesh size and leaf litter breakdown rates did not differ between ?ne and coarse mesh cages, suggesting that macroinvertebrates do not influence leaf litter decomposition in either habitat at our studied site. Leaf breakdown rates were faster in aquatic than in terrestrial habitats and the magnitude of these differences were comparable to studies in temperate regions, suggesting that equivalent factors can influence between‐habitat differences detected in our study.     

Conservation and restoration of New Zealand Island ecosystems

An ecological collapse has precipitated pioneering conservation initiatives in New Zealand. Many terrestrial communities in t he New Zealand archipelago have been devastated by over-exploitation, introduced mammals and habitat destruction. More recently, marine ecosystems have been depleted by over-harvesting. To mitigate against these losses, conservation in terrestrial environments has focused on protection of species and habitats. A similar approach is now under way in marine environments with the establishment of ‘no-take’ marine reserves. On land, conservation is now reaching beyond protection t o the eradication of pests from islands and restoration of their terrestrial ecosystems. Restoration on islands not only reduces threats to rare species; it also raises opportunities to investigate how species interact. In the sea, marine reserves not only enhance the diversity of depleted marine communities; they may also augment stocks of commercially harvested species. These initiatives provide many lessons that could be applied to degraded habitats elsewhere.     

Vertical stratification influences global patterns of biodiversity

Species distributions in terrestrial ecosystems are three‐dimensional, spanning both the horizontal landscape and the vertical space provided by the physical environment. Classical hypotheses suggest that communities become more vertically stratified with increasing species richness, owing to reduced competition or finer niche subdivision. However, this assertion remains untested in the context of the broader realm of biogeography. Here, integrating traits and distribution data for amphibians globally, we show how vertical strategies interact with the physical and climatic environments to govern global patterns of species richness and community composition. Our results reveal a marked latitudinal shift in strategies of vertical habitat use, from highly arboreal assemblages in the tropics to highly fossorial assemblages in sub‐tropical and temperate regions. Arboreality is strongly associated with precipitation, vegetation structure and climatic stability, whereas fossoriality is more common in dry environments with high diurnal temperature range and low vegetation structure. These analyses shed light on the importance of vertical stratification for species coexistence in species‐rich regions. As certain tropical habitats become drier from climate change, the rich biological diversity that is emblematic of the tropics may transition from vertically stratified to ‘flattened’, with future communities living mostly on or beneath the ground.     

Biogeography,habitat transitions and hybridization in a radiation of South American silverside fishes revealed by mitochondrial and genomic RAD data

Rivers and lake systems in the southern cone of South America have been widely influenced by historical glaciations, carrying important implications for the evolution of aquatic organisms, including prompting transitions between marine and freshwater habitats and by triggering hybridization among incipient species via waterway connectivity and stream capture events. Silverside fishes (Odontesthes) in the region comprise a radiation of 19 marine and freshwater species that have been hypothesized on the basis of morphological or mitochondrial DNA data to have either transitioned repeatedly into continental waters from the sea or colonized marine habitats following freshwater diversification. New double digest restriction‐site associated DNA data presented here provide a robust framework to investigate the biogeographical history of and habitat transitions in Odontesthes. We show that Odontesthes silversides originally diversified in the Pacific but independently colonized the Atlantic three times, producing three independent marine‐to‐freshwater transitions. Our results also indicate recent introgression of marine mitochondrial haplotypes into two freshwater clades, with more recurring instances of hybridization among Atlantic‐ versus Pacific‐slope species. In Pacific freshwater drainages, hybridization with a marine species appears to be geographically isolated and may be related to glaciation events. Substantial structural differences of estuarine gradients between these two geographical areas may have influenced the frequency, intensity and evolutionary effects of hybridization events.     

Wintering space use and site fidelity in a nomadic species,the snowy owl

Migratory species can exploit many habitats over vast geographic areas and adopt various patterns of space and habitat use throughout their annual cycle. In nomadic species, determinants of habitat use during the non‐breeding season are poorly known due to the unpredictability of their movement patterns. Here, we analysed variability in wintering space and habitat use by a highly nomadic species, the snowy owl, in eastern North America. Using 21 females tracked by satellite telemetry between 2007 and 2016, we 1) assessed how space use patterns in winter varied according to the type of environment (marine vs terrestrial), latitudinal zone (Arctic vs temperate), local snow conditions and lemming densities and 2) investigated winter habitat and site fidelity. Our results confirmed a high inter‐individual variation in patterns of habitat use by wintering snowy owls. Highly‐used areas were concentrated in the Arctic and in the marine and coastal environments. Owls wintering in the marine environment travelled over longer distances during the winter, had larger home ranges and these were divided in more smaller zones than individuals in terrestrial environments. Wintering home range sizes decreased with high winter lemming densities, use of the marine environment increased following high summer lemming densities, and a thick snow cover in autumn led to later settlement on the wintering ground. Contrary to expectations, snowy owls tended to make greater use of the marine environment when snow cover was thin. Snowy owls were highly consistent in their use of a given wintering environment and a specific latitudinal zone between years, but demonstrated flexibility in their space use and a modest site fidelity. The snowy owls’ consistency in wintering habitat use may provide them with advantages in terms of experience but their mobility and flexibility may help them to cope with changing environmental conditions at fine spatial scale.     

再论中国植被分区的原则和方案

 本文首先提出中国植被分区的原则和依据以及高级分区单位的标志,将全国划分为八大植被区,其中有五个区包括两个亚区,因作者主张亚区与区作为同一级的辅助单位看待,所以实际上把全国分为13个高级植被分区单位。除少数例外,每一植被区或亚区都分为一过渡带和典型带。全文以植被区或亚区、植被地带或亚地带为单位,论述其植被特点。     

Ecological Diversity in South American Mammals: Their Geographical Distribution Shows Variable Associations with Phylogenetic Diversity and Does Not Follow the Latitudinal Richness Gradient

The extent to which the latitudinal gradient in species richness may be paralleled by a similar gradient of increasing functional or phylogenetic diversity is a matter of controversy. We evaluated whether taxonomic richness (TR) is informative in terms of ecological diversity (ED, an approximation to functional diversity) and phylogenetic diversity (AvPD) using data on 531 mammal species representing South American old autochthonous (marsupials, xenarthrans), mid-Cenozoic immigrants (hystricognaths, primates) and newcomers (carnivorans, artiodactyls). If closely related species are ecologically more similar than distantly related species, AvPD will be a strong predictor of ED; however, lower ED than predicted from AvPD may be due to species retaining most of their ancestral characters, suggesting niche conservatism. This pattern could occur in tropical rainforests for taxa of tropical affinity (old autochthonous and mid-Cenozoic immigrants) and in open and arid habitats for newcomers. In contrast, higher ED than expected from AvPD could occur, possibly in association with niche evolution, in arid and open habitats for taxa of tropical affinity and in forested habitats for newcomers. We found that TR was a poor predictor of ED and AvPD. After controlling for TR, there was considerable variability in the extent to which AvPD accounted for ED. Taxa of tropical affinity did not support the prediction of ED deficit within tropical rainforests, rather, they showed a mosaic of regions with an excess of ED interspersed with zones of ED deficit within the tropics; newcomers showed ED deficit in arid and open regions. Some taxa of tropical affinity showed excess of ED in tropical desert areas (hystricognaths) or temperate semideserts (xenarthrans); newcomers showed excess of ED at cold-temperate latitudes in the Northern Hemisphere. This result suggests that extreme climatic conditions at both temperate and tropical latitudes may have promoted niche evolution in mammals.     

Latitudinal variation of diversity in European freshwater animals is not concordant across habitat types

Aim   We analysed the variation of species richness in the European freshwater fauna across latitude. In particular, we compared latitudinal patterns in species richness and β-diversity among species adapted to different habitat types.
Location   Europe.
Methods   We compiled data on occurrence for 14,020 animal species across 25 pre-defined biogeographical regions of European freshwaters from the Limnofauna Europaea . Furthermore, we extracted information on the habitat preferences of species. We assigned species to three habitat types: species adapted to groundwater, lotic (running water) and lentic (standing water) habitats. We analysed latitudinal patterns of species richness, the proportion of lentic species and β-diversity.
Results   Only lentic species showed a significant species–area relationship. We found a monotonic decline of species richness with latitude for groundwater and lotic habitats, but a hump-shaped relationship for lentic habitats. The proportion of lentic species increased from southern to northern latitudes. β-Diversity declined from groundwater to lentic habitats and from southern to northern latitudes.
Main conclusions   The differences in the latitudinal variation of species richness among species adapted to different habitat types are in part due to differences in the propensity for dispersal. Since lentic habitats are less persistent than lotic or groundwater habitats, lentic species evolved more efficient strategies for dispersal. The dispersal propensity of lentic species facilitated the recolonization of central Europe after the last glaciation. Overall, we stress the importance of considering the history of regions and lineages as well as the ecological traits of species for understanding patterns of biodiversity.     

EVOLUTIONARY AND BIOGEOGRAPHIC ORIGINS OF HIGH TROPICAL DIVERSITY IN OLD WORLD FROGS (RANIDAE)

Differences in species richness between regions are ultimately explained by patterns of speciation, extinction, and biogeographic dispersal. Yet, few studies have considered the role of all three processes in generating the high biodiversity of tropical regions. A recent study of a speciose group of predominately New World frogs (Hylidae) showed that their low diversity in temperate regions was associated with relatively recent colonization of these regions, rather than latitudinal differences in diversification rates (rates of speciation–extinction). Here, we perform parallel analyses on the most species-rich group of Old World frogs (Ranidae; ∼1300 species) to determine if similar processes drive the latitudinal diversity gradient. We estimate a time-calibrated phylogeny for 390 ranid species and use this phylogeny to analyze patterns of biogeography and diversification rates. As in hylids, we find a strong relationship between the timing of colonization of each region and its current diversity, with recent colonization of temperate regions from tropical regions. Diversification rates are similar in tropical and temperate clades, suggesting that neither accelerated tropical speciation rates nor greater temperate extinction rates explain high tropical diversity in this group. Instead, these results show the importance of historical biogeography in explaining high species richness in both the New World and Old World tropics.     

Multiple invasions into freshwater by pufferfishes (teleostei: tetraodontidae): a mitogenomic perspective

Pufferfishes of the Family Tetraodontidae are the most speciose group in the Order Tetraodontiformes and mainly inhabit coastal waters along continents. Although no members of other tetraodontiform families have fully discarded their marine lives, approximately 30 tetraodontid species spend their entire lives in freshwaters in disjunct tropical regions of South America, Central Africa, and Southeast Asia. To investigate the interrelationships of tetraodontid pufferfishes and thereby elucidate the evolutionary origins of their freshwater habitats, we performed phylogenetic analysis based on whole mitochondrial genome sequences from 50 tetraodontid species and closely related species (including 31 newly determined sequences). The resulting phylogenies reveal that the family is composed of four major lineages and that freshwater species from the different continents are independently nested in two of the four lineages. A monophyletic origin of the use of freshwater habitats was statistically rejected, and ancestral habitat reconstruction on the resulting tree demonstrates that tetraodontids independently entered freshwater habitats in different continents at least three times. Relaxed molecular-clock Bayesian divergence time estimation suggests that the timing of these invasions differs between continents, occurring at 0-10 million years ago (MA) in South America, 17-38 MA in Central Africa, and 48-78 MA in Southeast Asia. These timings are congruent with geological events that could facilitate adaptation to freshwater habitats in each continent.