Biogeography of Iberian freshwater fishes revisited: the roles of historical versus contemporary constraints

Aim The question of how much of the shared geographical distribution of biota is due to environmental vs. historical constraints remains unanswered. The aim of this paper is to disentangle the contribution of historical vs. contemporary factors to the distribution of freshwater fish species. In addition, it illustrates how quantifying the contribution of each type of factor improves the classification of biogeographical provinces. Location Iberian Peninsula, south‐western Europe (c. 581,000 km²). Methods We used the most comprehensive data on native fish distributions for the Iberian Peninsula, compiled from Portuguese and Spanish sources on a 20‐km grid‐cell resolution. Overall, 58 species were analysed after being categorized into three groups according to their ability to disperse through saltwater: (1) species strictly intolerant of saltwater (primary species); (2) species partially tolerant of saltwater, making limited incursions into saltwaters (secondary species); and (3) saltwater‐tolerant species that migrate back and forth from sea to freshwaters or have invaded freshwaters recently (peripheral species). Distance‐based multivariate analyses were used to test the role of historical (basin formation) vs. contemporary environmental (climate) conditions in explaining current patterns of native fish assemblage composition. Cluster analyses were performed to explore species co‐occurrence patterns and redefine biogeographical provinces based on the distributions of fishes. Results River basin boundaries were better at segregating species composition for all species groups than contemporary climate variables. This historical signal was especially evident for primary and secondary freshwater fishes. Eleven biogeographical provinces were delineated. Basins flowing to the Atlantic Ocean north of the Tagus Basin and those flowing to the Mediterranean Sea north of the Mijares Basin were the most dissimilar group. Primary and secondary freshwater species had higher province fidelity than peripheral species. Main conclusions The results support the hypothesis that historical factors exert greater constraints on native freshwater fish assemblages in the Iberian Peninsula than do current environmental factors. After examining patterns of assemblage variation across space, as evidenced by the biogeographical provinces, we discuss the likely dispersal and speciation events that underlie these patterns.     

Regional and longitudinal patterns of fish community structure in the Seine River basin, France

The fish communities of 371 sites from 4 natural regions of the Seine River basin were studied. The sites were located from small to medium size rivers (catchment area : 5 to 3895 km²). We examined the differences between local communities according to river size (estimated by catchment area) and region. In the Seine River basin, fish communities follow a general organisation rule: total species richness increases with river size and importance of limnophilic species versus rheophilic ones increases from upstream to downstream. However, fish communities show differences of total species richness, species richness of reproductive groups and species composition between the four natural regions of the basin. Particularly, river size and regional organisation of environmental factors interact on species composition of communities and several regional patterns of longitudinal changes of fish communities are identified. The origin and range of regional differences of fish communities are discussed according to historical and environmental factors.     

Biogeographical patterns of freshwater micro‐ and macroorganisms: a comparison between phytoplankton,zooplankton and fish in the eastern Mediterranean

Aim We investigated the biogeographical patterns of phytoplankton, zooplankton and fish in freshwater ecosystems. We tested whether spatial distance or environmental heterogeneity act as potential factors controlling community composition. Location Northern and central Greece, eastern Mediterranean. Method Data on 310 phytoplankton, 72 zooplankton and 37 fish species were collected from seven freshwater systems. Species occurrence data were used to generate similarity matrices describing community composition. We performed Mantel tests to compare spatial patterns in community composition of phytoplankton, zooplankton and fish. Next, we examined the correlation between geographical distance and the degree of similarity in community composition. The analysis was repeated for different taxonomic, trophic and size‐based groups of the microorganisms studied. We assessed different environmental variables (topographic and limnological) as predictors of community composition. Results Phytoplankton community composition showed a strong positive correlation with environmental heterogeneity but was not correlated with the geographical distance between systems. Zooplankton community composition was unrelated to geographical distance and was only weakly correlated with environmental variables. In contrast, fish community similarity decayed significantly with distance. We found no relationship along all pairwise comparisons of the compositional matrices of the three groups. The pairwise comparisons of the different taxonomic, trophic and size‐based groups of the microorganism communities studied were in accordance with the results for the entire microorganism community. Main conclusions Our results support the proposition that the biogeography of microorganisms does not demonstrate a distance–decay pattern and further suggest that, in reality, the drivers of distribution depend on the specific community examined. In contrast, the biogeography of macroorganisms was affected by geographical distance. These differences reflect the dispersal abilities of the different organisms. The microorganisms exhibit passive dispersal through the air, with local environmental conditions structuring their community composition. On the other hand, for macroorganisms such as fish, the terrestrial environment could pose barriers to their dispersal; with fish structuring distinctive communities over greater distances. Overall, we suggest that the biogeography of freshwater phytoplankton and zooplankton reflects contemporary environmental conditions, while the biogeographical patterns for fish inhabiting the same systems are related to factors affecting their dispersal ability.     

Darwin, historical biogeography, and the importance of overcoming binary opposites

It is well known that Darwin and Wallace came to discover the phenomenon of evolution through a historical approach to the geographical distribution of organisms. Before Darwin, evolution was a mere speculation that could be invoked to explain some facts. Darwin's biogeographical argument for evolution is based largely on three main explanatory hypotheses. The first is that the geographical distribution of organisms is historically informative. The second hypothesis is that long-distance dispersal over barriers is one main force (extinction is the other) that modifies the distribution of organisms. The third of Darwin's biogeographical hypotheses is that the factors that shape the distribution of organisms are mainly historical (large, often global and long temporal scales) rather than ecological (small spatial and short temporal scales). From the time of Darwin until now, a wide spectrum of biogeographical schools have provided new insights that challenge the central role of space, dispersal and history as the main explanatory hypotheses for the distribution of organisms, generating three binary opposites: (1) the spatial dimension of evolution: geographical distribution of organisms as historically informative vs. historically uninformative; (2) the processes that modify the geographical distribution of organisms: dispersal vs. vicariance; and (3) the explanation of geographical distribution: history vs. ecology. We analyse these three binary opposites to show that the components of each are complementary rather than antagonistic approaches to the study of biogeography.     

Three ecological factors influencing riverine fish diversity in the Shubuto River system,Japan: habitat capacity,habitat heterogeneity and immigration

A major goal of stream ecology is to identify environmental gradients that shape riverine communities. We examined the relative importance of three ecological factors that have been hypothesized to influence a longitudinal pattern of fish diversity: habitat capacity, heterogeneity and immigration of diadromous fishes. Field surveys were carried out in the entire network of the Shubuto River system, Hokkaido, Japan. A hierarchical partitioning approach revealed that distance from the sea, a proxy for immigration potential of diadromous fishes, had the greatest explanatory capacity, by which 24.9 % of variation in fish species richness was explained. Habitat capacity (approximated by catchment area) was also identified as a significant predictor of fish diversity, whereas habitat heterogeneity brought little improvement to the model performance. These results reflect the fish fauna of the Shubuto River system, in which diadromous fishes are dominant in both abundance and species richness.     

The Ethiopian subregion fish fauna: an original patchwork with various origins

In this article, I analysed the biogeographical distribution and the historical origin of freshwater fish of the Ethiopian Rift Valley (ERV) region which is not a homogeneous area. The Ethiopian region, including Eritrea and part of Somalia, includes 13 basins as geographical units. A comparison was carried out with four Near East geographical units. A correspondence analysis was applied to presence–absence data for the 17 basins × 170 species’ matrix. Then, a faunistic distance (Euclidian distance) between basins was computed from the coordinates of factors retained in this new analysis. From this faunistic distance, a hierarchical cluster analysis was performed and a dendrogram produced by a UGPMA classification. The old Abyssinian ichthyo-province (sensu Roberts, Zool J Linn Soc 57:249–319, 1975) was not homogeneous in species composition. I distinguished four main groups based on the present species composition which seemed to be mainly influenced by historical events. The species composition of the ERV basins was not randomly distributed and can be classified into four main geographical units. The importance of the historical events was demonstrated and two origins can be considered. The westernmost basins are under Nilo-Sudan influence whereas the easternmost ones seem to have been colonised from the Near East. In this Eastern part it can be distinguished three different groups: the main ERV province and two derived sub-regions, Lake Tana with a Labeobarbus intermedius related species flock and the Coastal Red Sea, characterised by a very impoverished freshwater fish fauna.     

Environmental factors determining the structure of rotifer communities in a river-shed reservoir

This study was designed to find out which environmental factors (temperature, oxygen concentration and food resources) are more important in steering the rotifers vertical distribution and community structure in an oligo-changing to-mesotrophic reservoir. It was observed that, despite the influence of temperature and oxygen content of the water, biotic factors especially the food availability and changes in trophic state were crucial in shaping the vertical distribution and species composition of rotifers in Meimoa Reservoir, located in Tagus River in Portugal.     

Composition, temporal changes and ecological guild classification of the ichthyofaunas of large European estuaries – a comparison between the Tagus (Portugal) and the Elbe (Germany)

Fish species densities from 1025 hauls obtained with beam trawl and stow net catches from the southern Tagus and northern Elbe estuaries were analysed for two different periods of time between 1978 and 1997. The data were related with environmental factors to compare species composition and densities, contributions of different ecological guilds, structuring environmental key factors, intra‐annual cyclicity and interperiodical stability of ichthyofaunas in large European estuaries at different latitudes. Although the total number of species in both the Tagus and Elbe estuaries (Tagus: 57 species, Elbe: 58 species) was very similar, significant differences were estimated in species compositions, species densities and number of species, and individuals of the different ecological guilds between both estuaries. A total of 15 species occurred in both estuaries but only two of these species, the catadromous Anguilla anguilla and the marine estuarine opportunist Pleuronectes flesus, contributed substantially to the total abundance. Marine species together contributed more than 5% of the total number of species in the Tagus and Elbe estuaries, whereas their contribution in number of individuals amounted to 22% in the Tagus and only 9% in the Elbe. In terms of number of individuals, the ichthyofauna of the Tagus estuary was dominated by estuarine species (especially Pomatoschistus spp.) contributing 68% of the total abundance. In contrast, the fish fauna of the Elbe estuary was dominated by high numbers of anadromous species (88%) individuals, especially Osmerus eperlanus. Canonical corresponding analyses (CCA) indicated that, besides intra‐annual variables, salinity and mouth distance were two of the most important environmental factors structuring the fish communities in both estuaries. Moreover, water depth was found to be an important environmental factor in the Tagus estuary, whereas the water temperature substantially affected the ichthyofaunal composition in the Elbe estuary. In both estuaries, contributions by number of species and number of individuals of the different ecological guilds were not statistically different between the two periods considered, 1978–80 and 1995–97 for the Tagus, and 1981–86 and 1989–95 for the Elbe. Nevertheless, from 1981–86 until 1989–95 a remarkable increase of about 4.5 times in the densities of O. eperlanus was observed in the Elbe estuary, probably related to improved water quality, especially of oxygen conditions in the nurseries since 1990. This was also supported by the results of CCA, suggesting important influences of the years 1989 and 1990 on the Elbe estuary fish assemblage. Climate change as a result of global warming may be evident for the observed changes of the densities of P. flesus which decreased in the southern Tagus estuary and, in contrast, increased in the northern Elbe estuary during comparable periods of time. It is recommended to use the ecological guild classification scheme which was applied for the Tagus and Elbe estuaries for further studies of European estuarine fish assemblages.     

Faunal breaks and species composition of Indo-Pacific corals: the role of plate tectonics,environment and habitat distribution

Species richness gradients are ubiquitous in nature, but the mechanisms that generate and maintain these patterns at macroecological scales remain unresolved. We use a new approach that focuses on overlapping geographical ranges of species to reveal that Indo-Pacific corals are assembled within 11 distinct faunal provinces. Province limits are characterized by co-occurrence of multiple species range boundaries. Unexpectedly, these faunal breaks are poorly predicted by contemporary environmental conditions and the present-day distribution of habitat. Instead, faunal breaks show striking concordance with geological features (tectonic plates and mantle plume tracks). The depth range over which a species occurs, its larval development rate and genus age are important determinants of the likelihood that species will straddle faunal breaks. Our findings indicate that historical processes, habitat heterogeneity and species colonization ability account for more of the present-day biogeographical patterns of corals than explanations based on the contemporary distribution of reefs or environmental conditions.     

Evolutionary history of the fish genus <Emphasis Type="Italic">Astyanax</Emphasis> Baird &; Girard (1854) (Actinopterygii,Characidae) in Mesoamerica reveals multiple morphological homoplasies

Background  

Mesoamerica is one of the world's most complex biogeographical regions, mostly due to its complex geological history. This complexity has led to interesting biogeographical processes that have resulted in the current diversity and distribution of fauna in the region. The fish genus Astyanax represents a useful model to assess biogeographical hypotheses due to it being one of the most diverse and widely distributed freshwater fish species in the New World. We used mitochondrial and nuclear DNA to evaluate phylogenetic relationships within the genus in Mesoamerica, and to develop historical biogeographical hypotheses to explain its current distribution.     

柳江鱼类群落结构及多样性研究

为了解柳江鱼类群落结构、多样性及其与环境因子的关系, 2018年7月和12月在柳江9个江段(榕江、从江、融安、柳城、象州、永福、鹿寨、河池和宜州)开展鱼类和环境调查。共采集鱼类36474尾, 隶属于5目20科80属115种; 其中, 鲤科鱼类最多(68种), 占59.13%。Shannon-Wiener多样性指数、Simpson指数、Pielou均匀度指数和Margalef丰富度指数的变化范围分别为2.13—3.12、0.81—0.93、0.58—0.80和4.65—7.18。鱼类优势种以银Squalidus argentatus、大眼华鳊Sinibrama macrops、胡Microphysogobio chenhsienensis、银鲴Xenocypris macrolepis和南方拟?Pseudohemiculter dispar等小型鱼类为主。丰度/生物量比较曲线表明, 宜州、榕江、鹿寨、柳城、象州和融安江段鱼类群落受到中度或严重干扰, 河池、从江和永福鱼类群落结构相对稳定。采用非度量多维标度排序(NMDS)方法对鱼类群落的时空分布特征进行分析, 将鱼类划分为3个类群: 柳江上游的从江-榕江类群, 洛清江支流的永福-鹿寨类群, 柳江中下游及龙江支流的融安-柳城-象州-河池-宜州类群。采用典范对应分析(CCA)分析鱼类群落与环境因子的关系, 发现河宽、海拔和pH是柳江鱼类群落结构差异的主要影响因子。与历史资料相比, 柳江鱼类群落结构发生显著变化, 经济鱼类占比减少, 鱼类个体呈现小型化和低龄化。建议从以下几个方面加强柳江鱼类多样性保护: 加强执法力度, 严厉打击偷捕、电捕等非法行为; 对主要经济鱼类开展增殖放流; 恢复河流的连通性; 对流域珍稀特有鱼类进行监测、救护和繁育。     

珠江水系鱼类群落多样性空间分布格局

珠江是我国南方第一大河,是我国重要淡水渔业生产基地和水生生物资源基因库。珠江鱼类在维持生物多样性、提供鱼类种质资源方面举足轻重。但是到目前为止,关于其鱼类空间分布格局的研究甚少。特别是近几十年来各种水工建设和过度捕捞使得渔业资源急剧衰退,鱼类空间分布的研究显得尤为重要。2015年对珠江全流域13个站位进行了全面调查,共采集渔获物10119尾,隶属于94种72属17科。鲤科鱼类占显著优势,其次种类较多的依次为鲿科、鳅科。采用非度量多维标度排序(NMDS)方法对鱼类群落空间分布特征进行了分析,结果表明珠江鱼类被划分为3个类群,即以餐、南方拟餐、黄颡鱼等小型鱼类为主的中上游类群、以赤眼鳟、鲮鱼、广东鲂等中型鱼类为主的中下游类群和以罗非鱼为主的重要支流类群。同时发现中下游物种多样性高,上游及河口江段多样性低的格局。采用冗余分析方法(RDA)分析了鱼类多样性与环境因子的关系,发现年均气温、降雨量、年均径流量、河流宽度与透明度是珠江水系河流鱼类群落结构差异的主要影响因子,其中年均气温是影响鱼类群落分布的最关键因子之一。与历史资料对比后发现,珠江鱼类种类明显减少、空间分布也发生了巨大改变。研究是珠江水系野生渔业资源长期调查的一部分,研究结果将对渔业资源的多样性保护和可持续利用具有指导意义。     

Global patterns and predictors of tropical reef fish species richness

In the marine realm, the tropics host an extraordinary diversity of taxa but the drivers underlying the global distribution of marine organisms are still under scrutiny and we still lack an accurate global predictive model. Using a spatial database for 6336 tropical reef fishes, we attempted to predict species richness according to geometric, biogeographical and environmental explanatory variables. In particular, we aimed to evaluate and disentangle the predictive performances of temperature, habitat area, connectivity, mid‐domain effect and biogeographical region on reef fish species richness. We used boosted regression trees, a flexible machine‐learning technique, to build our predictive model and structural equation modeling to test for potential ‘mediation effects’ among predictors. Our model proved to be accurate, explaining 80% of the total deviance in fish richness using a cross‐validated procedure. Coral reef area and biogeographical region were the primary predictors of reef fish species richness, followed by coast length, connectivity, mid‐domain effect and sea surface temperature, with interactions between the region and other predictors. Important indirect effects of water temperature on reef fish richness, mediated by coral reef area, were also identified. The relationship between environmental predictors and species richness varied markedly among biogeographical regions. Our analysis revealed that a few easily accessible variables can accurately predict reef fish species richness. They also highlight concerns regarding ongoing environmental declines, with region‐specific responses to variation in environmental conditions predicting a variable response to anthropogenic impacts.     

Using ground‐derived data to assess the environmental niche of the spinose ear tick,Otobius megnini

Four layers of environmental information, namely 1) monthly atmospheric temperature and rainfall, 2) annual ground and underground moisture, evaporation, surface adjacent specific humidity, and temperature, 3) monthly Normalized Derived Vegetation Index (NDVI), and 4) soil physical attributes, were used separately to define the expected geographical distribution and environmental niche of the spinose ear tick, Otobius megnini (Dugès) (Acari: Ixodida: Argasidae), an endophilic argasid, in both tropical and neotropical regions. The best predictive values were obtained from ground‐derived climate. Air‐derived features ranked second. The remaining environmental information had poor discriminatory abilities. The most informative variables in the distribution of neotropical populations are ground temperatures, with surface humidity ranking second. In the tropics, surface humidity is the most important factor delineating the distribution of O. megnini. Marginality scores are similar for tick populations in both biogeographical regions, but specialization factors are different, supporting the findings that both populations are regulated by different variables. Similarly, models trained with records of one biogeographical region and projected into the other one, resulted in poorer predictions than when trained with the homologous set of records. Populations of the tick in the tropics experience a different range of temperatures than their neotropical relatives, whereas marginality scores are similar. The conclusion is that each population uses particular portions of the environmental niche, probably because of different climate or competitor constraints on either biogeographical region.     

Biogeographical regions of the Iberian peninsula based on freshwater fish and amphibian distributions

We classified the main Iberian river basins based on the presence and absence of freshwater fishes and amphibians. For both taxonomic groups we analysed three data sets; 1) endemic species only, to search for biotic boundaries related to historical events, 2) indigenous species, which include endemic ones, to search for biotic boundaries related to ecological factors, 3) indigenous and well-established introduced species, to assess the influence of man in the current biogeographical patterns of fishes and amphibians. We used both phenetic and cladistic methods, followed by a consensus analysis to provide an overall biogeographical pattern. Based on all fish distributions, the Iberian Peninsula is divided into three biogeographical regions: Cantabrian, Atlantic and Mediterranean, No boundary existed between the Cantabrian and Atlantic regions when only indigenous fish species were considered. This suggests that this boundary has been induced by man, probably through the differential introduction of fish species into reservoirs at one or other side of the boundary. Run-off and the size of the river basins are the environmental factors that distinguished the Atlantic and Mediterranean regions. However, regionalization based only on endemic freshwater fishes showed a latitudinal pattern that agrees with the paleogeographic events of the Upper Oligocene-Lower Miocene period. By contrast, one northern and one southern region were distinguished based on all amphibian distributions and on indigenous amphibians only, which suggests that human activity has not significantly affected the overall biogeographical pattern of amphibians in the Iberian Peninsula. Interannual predictability of precipitation best accounts for this regionalization. Based on endemic amphibians, the Iberian Peninsula is divided into three regions that closely resemble the three separate land areas of the Upper Eocene-Lower Oligocene period. The consensus between the biogeographical regions based on fishes and amphibians yields five pairs of basins. Geological origin of the basins seems to better explain the consensus between the biogeographical patterns of fishes and amphibians, whereas ecological factors probably contribute to the differences between them.     

Historical biogeography of South American freshwater fishes

Aim To investigate biogeographical patterns of the obligate freshwater fish order Characiformes. Location South America. Methods Parsimony analysis of endemicity, likelihood analysis of congruent geographical distribution, and partition Bremer support were used. Results Areas of endemism are deduced from parsimony analysis of endemicity, and putative dispersal routes from a separate analysis of discordant patterns of distribution. Main conclusions Our results demonstrate the occurrence of 11 major areas of endemism and support a preferential eastern–western differentiation of the characiforms in the Amazonian region, contrasting with the southern–northern differentiation of terrestrial organisms. The areas of endemism identified seem to be deeply influenced by the distribution of the emerged land during the 100‐m marine highstand that occurred during the late Miocene and allow us to hypothesize the existence of eight aquatic freshwater refuges at that time. The raw distribution of non‐endemic species supports nine patterns of species distribution across the 11 areas of endemism, two of which support a southern–northern differentiation in the eastern part of the Amazon. This result shows that the main channel of the Amazon limited dispersal between tributaries from each bank of the river. The levels of endemism further demonstrate that the aquatic freshwater refuges promoted allopatric speciation and later allowed the colonization of the lowlands. By contrast, the biogeographical pattern found in the western part of the Amazon is identified as a result of the Miocene Andean foreland dynamic and the uplift of the palaeoarches that promoted allopatric divergence across several sedimentary basins by the establishment of disconnected floodplains. The assessment of conflicting species distributions also shows the presence of seven putative dispersal routes between the Amazon, Orinoco and Paraná rivers. Our findings suggest that, rather than there being a single predominant process, the establishment of the modern South American freshwater fish biotas is the result of an interaction between marine incursions, uplift of the palaeoarches, and historical connections allowing cross‐drainage dispersal.     

Determinants of the geographical extent of invasive plants in China: effects of biogeographical origin,life cycle and time since introduction

Plant invasions cause serious impacts to environment and human economy, and lots of researches have been paid to examine factors that determine the geographical extent of plant invaders. Many intrinsic species attributes, external environmental factors, and time since introduction have all been proposed to explain invasive success. Data on invasive plants distribution in province regions in China were collected, and species attributes and historical factors such as biogeographical origin, life cycle, and time since introduction were determined. While some annuals may eventually occupy the whole country area, perennials are unlikely to do so, particularly those from Central and South America. Time since introduction is significantly related to the number of provinces occupied by an invader; nevertheless after control of certain residence time, annuals are not likely to more provinces than the perennial herbs. High-impact invaders tend to spread faster than other invaders, but only when their life cycle is taken into account. This suggests that other plant traits, instead of life cycle, along with extrinsic factors, greatly contribute to invasive success and distribution area. Invasive plants in China are mainly distributed in human-made habitats, and their long-distance dispersal is primarily associated with human activities. Future studies need to consider interactions between species traits, environmental conditions, and human activities for further insights in researches alike.     

Population genetics of a marine bivalve, Pinctada maxima, throughout the Indo-Australian Archipelago shows differentiation and decreased diversity at range limits

Intraspecific genetic diversity governs the potential of species to prevail in the face of environmental or ecological challenges; therefore, its protection is critical. The Indo-Australian Archipelago (IAA) is a significant reservoir of the world's marine biodiversity and a region of high conservation priority. Yet, despite indications that the IAA may harbour greater intraspecific variation, multiple-locus genetic diversity data are limited. We investigated microsatellite DNA variation in Pinctada maxima populations from the IAA to elucidate potential factors influencing levels of genetic diversity in the region. Results indicate that genetic diversity decreases as the geographical distance away from central Indonesia increases, and that populations located towards the centre of P. maxima 's range are more genetically diverse than those located peripherally ( P <  0.01). Significant partitioning of genetic variation was identified ( F _ST = 0.027; R _ST = 0.023, P  < 0.001) and indicates that historical biogeographical episodes or oceanographic factors have shaped present population genetic structure. We propose that the genetic diversity peak in P. maxima populations may be due to (i) an abundance of suitable habitat within the IAA, meaning larger, more temporally stable populations can be maintained and are less likely to encounter genetic bottlenecks; and/or (ii) the close proximity of biogeographical barriers around central Indonesia results in increased genetic diversity in the region because of admixture of genetically divergent populations. We encourage further genetic diversity studies of IAA marine biota to confirm whether this region has a significant role in maintaining intraspecific diversity, which will greatly assist the planning and efficacy of future conservation efforts.     

The other face of Lyell: historical biogeography in his Principles of geology

Although some excellent articles about Lyell's work have been published, they do not explicitly deal with Lyell's biogeographical conceptions. The purpose of this paper is to analyse Lyell's biogeographical model in terms of its own internal structure. Lyell tried to explain the distribution of organisms by appealing to a real cause (climate). However, he was aware that environmental conditions were clearly insufficient to explain the existence of biogeographical regions. Lyell's adherence to ecological determinism generated strong tensions within his biogeographical model. He shifted from granting a secondary weight to dispersal to assigning it a major role. By doing so, Lyell was led into an evident contradiction. A permanent tension in Lyell's ideas was generated by the prevalent explanatory pattern of his time. The explanatory model based on laws did not produce satisfactory results in biology because it did not deal with historical processes. We may conclude that the knowledge of organic distribution interested Lyell as long as it could be explained by the uniformitarian principles of his geological system. The importance of the second volume of the Principles of geology lies in its ample and systematic argumentation about the geographical distribution of organisms. Lyell established, independently from any theory about organic change, the first version of dispersalist biogeography.     

Biogeographical indices of the freshwater fish spatial distribution in the upper reaches of the Yangtze River

The spatial distribution of freshwater fish species (Total species and Unique species) in the upper reaches of the Yangtze River was studied, focusing on the spatial distribution of geographic ranges. Two biogeographical indices: environmental resistance (R₅₀), anisotropy (A₅₀) and a fractal dimension method were used to identify the relationship between them and spatial distribution of freshwater fish. R₅₀ quantifies the loss of biotic resemblance occurring from any point in the map to the rest of the study area. A₅₀ quantifies the extent to which the perimeter: area ratio of the geographical ranges of all species whose distributions overlap at any particular location depart from the perimeter of area ratio of a circle. We concluded that the western of the study area is populated by a large number of total and unique species, the eastern part with the total species richness, the middle regions populated by small number of unique species, the mid-west region populated by small number of total species, and the mid-eastern region populated by a large number of total species. We also analyzed how topology affects the spatial arrangement of species, species density has strong curvilinear correlation (species density increases with increased of fractal dimension), the species density in the tributaries is lower than that in mainstream. The results and methods used give us detailed information about the spatial distribution of species, so as to illuminate the species distribution and change by the human activities.