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How many species are there, globally? Range of estimates: 2 – 100 million Best estimate: 10 million 1.4 – 2 million species have a name. An estimated 97% of all species are invertebrates, and within these, insects are by far the most numerous (an estimated 1-30 million species). In one study of just 19 trees in Panama, 1200 species of beetle were discovered , of which 80% were previously unknown to science. Within the vertebrates species: 23,500 ray-finned fish, 9000-10,000 birds, 7,984 reptiles, 5400 amphibians, 4475-5000 mammals. 70% of the world's species occur in only 12 countries: Australia, Brazil, China, Columbia, Ecuador, India, Indonesia, Madagascar, Mexico, Peru, and Zaire. On average, 3 new species of birds are found each year An estimated 40 percent of freshwater fishes in South America have not yet been classified. The deep sea floor may contain as many as a million undescribed species. Hydrothermal vent communities, discovered less than two decades ago, >20 new families or subfamilies, 50 new genera, and 100 new species were found. How to measure species diversity? Counting the number of species has several challenges: • species recognition (especially the small ones) • the influence of sampling effort and sample size 1 transect: 3 species 2 transects: 5 species 3 transects: 8 species 4 transects: 8 species 5 transects: 8 species The sampling artifact: Number of species in sample A bigger sample usually contains more species. Community 1 Community 2 Number of individuals in sample The sampling artifact: Fisher’s a Can be avoided by using diversity indices, rather than species numbers. community 1 community 2 Number of individuals in sample Fisher’s a does not change with sample size. How do you figure out how species numbers increase with area? Nested sampling design: 403 Total species number in an area: 403 species. Nested sampling design: 4 patches 280 288 312 341 Average species number of 305. How do you figure out of species numbers increase with area? Nested sampling design: 8 patches 220 269 256 301 245 242 305 Average species number of 266. 289 How do you figure out of species numbers increase with area? Nested sampling design: 40 patches 150 191 228 145 202 213 187 Average species number of 188. How do you figure out of species numbers increase with area? Nested sampling design: 150 patches 321 225 193 188 96 154 105 55 113 Average species number of 160. 150 450 Species Number 400 350 300 250 200 150 100 50 0 0 200 400 600 800 1000 1200 Log Species Number Area size (ha) 6.5 6 5.5 5 4.5 4 0 1 2 3 4 Log Area 5 6 7 8 Log Species Number The increase in the log of species number is proportional to the increase in log area. 6.5 6 5.5 5 4.5 4 0 1 2 3 4 Log Area 5 6 7 log S z log A log c S cA z 8 Examples: From Rosenzweig 1995 Log species number Generalized species-area curve for nested samples within continents: log S z log A log c Log area • z-values are quite similar between taxa, continents, ecosystems. • z-values are often in the range: 0.1-0.2. • c-values vary, expressing systematic differences in biodiversity between taxa, ecosystems, etc. Spatial patterns of biodiversity: Productivity gradient: More productive ecosystems usually have more species per area. But, highly productive ecosystems often have reduced diversity. Habitat heterogeneity gradient: More spatially variable environments have more species per area. Latitudinal gradient: Biodiversity decreases between the species-rich equatorial tropics and the species-poor polar regions. Island patterns: Islands have fewer species per area than their associated mainlands. The further away the island, the fewer species. Productivity gradient. Cold & dry Moist & warm Productivity From Rosenzweig 1995 Habitat heterogeneity gradient. Number of plants per 300 m2 plot beside the Hood River, Canada. From Rownsend, Begon and Harper 2003 Habitat heterogeneity gradient. Bird diversity in two continents From Rosenzweig 1995 Latitudinal gradient: From Rosenzweig 1995 Species-area curves for islands are different: Islands have fewer species than equal areas on the adjacent mainland. Species-area curves for islands are different: Islands have fewer species than equal areas on the adjacent mainland. islands mainland 42 55 160 141 188 266 305 180 403 Mainland: Islands: From Rosenzweig 1995 New Guinea: New Guinea islands: From Rosenzweig 1995 Species-area curves for islands are different: Island further from the mainland have fewer species than islands of equal size closer to the mainland. islands mainland 42 110 180 160 188 19 266 305 166 141 403 Log species number Generalized species-area curve for islands and their associated mainlands: log S z log S log c close far islands Log area • z-values for islands are usually greater than for mainlands. Generally, z = 0.2 – 0.6. • z-values are greater with the farther they are from the mainland. Interprovincial species-area curve: • larger continents (provinces) also have more species and the zvalue for interprovincial species-area curves is 1 or greater. Rosenzweig 1997 The three biological scales of species–area curves. Rosenzweig M L PNAS 2001;98:5404-5410 Temporal patterns of biodiversity: In succession: Diversity increases in the course of succession. With disturbance frequency: Intermediate disturbance frequencies have highest diversity. In evolutionary time: Over millions of years diversity stays relatively constant. That means, relatively fast recovery from mass extinction events. In the history of life on earth: Over hundreds of millions of years, diversity increased. Succession: Disturbance frequency: Disturbance From Rosenzweig 1995 Disturbance frequency: Algae and barnacles on rocks of different sizes From Rosenzweig 1995 Evolutionary time: Evolutionary time: From Rosenzweig 1995 Uintatherium Hyracotherium From Rosenzweig 1995 From Rosenzweig 1995 Triassic Devonian Ordovician Cambrian Pre-cambrian In the history of life on earth: Townsend, Begon and Harper 2003 Other patterns: Food web gradient: Higher trophic levels usually have lower diversity than lower trophic levels. Exception: there are more animal than plant species. There are more parasite than host species. Body size: Within a taxon, there are more intermediate-size species than either very large or very small species. Food web gradients: From Rosenzweig 1995 Patterns with body size Dragonflies and Damselflies From Rosenzweig 1995 Patterns with body size From Rosenzweig 1995 Patterns with body size From Rosenzweig 1995 Log species number Log area # species # species Latitude close far islands Log area # species Log species number Patterns in space productivity Habitat variety Species number Species number Time during succession Disturbance frequency Species number Species number Patterns in time Hundreds of Millions of years Millions of years # species # species Two more patterns Trophic level Body size within taxon