Synopsis of global fresh and brackish water occurrences of the bull shark Carcharhinus leucas Valenciennes, 1839 (Pisces: Carcharhinidae), with comments on distribution and habitat use

The bull shark (Carcharhinus leucas Valenciennes, 1839) is a large, primarily coastally distributed shark famous for its ability to penetrate far into freshwater bodies in tropical, subtropical, and warm-temperate climates. It is a cosmopolitan species with a geographical range that includes the coastlines of all major ocean basins (Atlantic Ocean, Indian Ocean, Pacific Ocean). As a consequence, freshwater occurrences of C. leucas are possible everywhere inside its geographic range. Carcharhinus leucas is a fully euryhaline, amphidromous species and possibly the widest-ranging of all freshwater tolerating elasmobranchs. This species is found not only in river systems with sea access that are not interrupted by human impediments but in hypersaline lakes as well. Rivers and estuaries are believed to be important nursery grounds for C. leucas, as suggested by observations of pregnant females in estuaries and neonates with umbilical scars in rivers and river mouths. Due to the physical capability of this species to enter riverine systems, the documentation of its occurrence in fresh and brackish water is essential for future conservation plans, fishery inspections, and scientific studies that focus on the link between low salinity habitats, shark nurseries, and feeding areas. The author's review of the available literature on C. leucas revealed the absence of a comprehensive overview of fresh and brackish water localities (rivers and associated lakes, estuaries) with C. leucas records. The purpose of this literature review is to provide a global list of rivers, river systems, lakes, estuaries, and lagoons with records and reports of this species, including a link to the used references as a base for regional, national, and international conservation strategies. Therefore, the objective of this work is to present lists of fresh and brackish water habitats with records of C. leucas as the result of an extensive literature review and analysis of databases. This survey also took into account estuaries and lagoons, regarding their function as important nursery grounds for C. leucas. The analysis of references included is not only from the scientific literature, but also includes semi-scientific references and the common press if reliable. The result of 415 global fresh and brackish water localities with evidence of C. leucas highlights the importance of these habitats for the reproduction of this species. Moreover, gaps in available distribution maps are critically discussed as well as interpretations and conclusions made regarding possible reasons for the distribution range of C. leucas, which can be interpreted as the result of geographic circumstances, but also as a result of the current state of knowledge about the distribution of this species. The results of the examination of available references were used to build a reliable and updated distribution map for C. leucas, which is also presented here.