Methods

Our first goal was to understand the relationship between primary productivity and species richness for several groups of freshwater organisms. By species richness, we mean the number of species observed in a lake over a number of years. It is useful to have several years of observations because the number of species observed varies from year to year. We chose the total list of species (the asymptote of the collectors curve) as our index of species richness. The lakes studied as part of the U.S. Long-Term Ecological Research (LTER) Program are particularly valuable because they have been studied for two decades, and complete species lists exist for many kinds of organisms in these systems. LTER lake sites occur in northern and southern Wisconsin and northern Alaska (Toolik Lake). However, because there are fewer than 15 LTER lakes (and only seven with measured rates of primary productivity), we increased sample size by including data from additional well-studied lakes of similar size, but which span a greater range of primary productivity (see Table 1). These lakes have been studied for several years, and estimates of annual primary productivity exist for each lake. Some well-studied lakes were not included, such as those which lacked much of the crucial data, or lakes that were unusually turbid or saline. For example, Lake Okeechobee (Florida, USA) is turbid and exhibits a wide range of productivity levels, depending on the part of the lake sampled, while Marion Lake (British Columbia, Canada) has a flushing rate of only a few days (W. E. Neill, personal communication). Sampling design and protocol are not standardized among studies of lakes. For example, species identifications were done by different people, sampling period was quite variable, and the number of samples per lake was variable. Such heterogeneity reduces the accuracy and precision of relationships between productivity and species richness.Primary productivity.—Pelagic primary productivity (PPR) can be measured by the 14C method (Vollenweider 1974). This method gives a close approximation to gross primary productivity (GPP), but because some of the fixed carbon is respired quickly, the value obtained is less than GPP (Fee et al.1982). Point values of PPR are then integrated by depth and area to produce estimates of whole-lake annual primary productivity per cubic meter or square meter.Lake primary productivity is fundamentally different than productivity measured in other biomes (e.g., grasslands, forests). The 14C method measures available (gross) primary productivity more than utilized (net) production, which is what is normally measured in terrestrial systems. The 14C method is also a fairly direct measure of productivity, compared to the proxy methods (e.g., nutrient loading, biomass, climate, soil fertility) used in many studies.Sampling protocols for aquatic organisms.—Sampling protocols differed among taxonomic groups and lakes (e.g., Downing and Rigler 1984). For example, phytoplankton samples are taken by capturing (at most) a few liters of lake water, either from a specific depth or with a sampler that integrates water across a range of depths. Zooplankton are usually sampled by vertical tows (i.e., raising a net through the water column). Both zooplankton and phytoplankton samples are typically taken from the center of the lake, although replicate samples at different locations may be taken from larger lakes. Planktonic organisms are much smaller than the sampling device, and hundreds to hundreds of thousands of organisms are typically captured in a single sample. In contrast, aquatic macrophytes are sampled using quadrats and rake samples, or simply based on a walk around the lake, while fish are sampled using a variety of nets andoror electroshocking equipment. Criteria for species lists.—Species lists for fish, macrophytes, and pelagic phytoplankton, rotifers, cladocerans, and copepods were obtained from the literature and from unpublished data. We avoided lists restricted to only dominant or common species, and thus included only lists that were exhaustive. Few lakes had species lists for all six groups of organisms. However, we included any lake that had an estimate of the average annual primary productivity and had lists for at least three taxa.We standardized this database by developing criteria for inclusion of species in analyses. Phytoplankton lists included all prokaryotic and eukaryotic photosynthetic phytoplankton for which there were abundances of more than one organism per milliliter (a criterion also used by Lewis 1979). We included all nonsessile species caught in open water as pelagic rotifers. For the crustacean zooplankton (cladocerans and copepods), we followed the criteria of Dodson (1992). Species lists of macrophytes included all submerged, floating, or emergent species of flowering plants, including Typha, sedges, grasses, and duck weed. We did not include Isoetes or macroalgae such as Chara and Nitella as macrophytes. The fish list included all species reported from the lake, including introduced taxa. Fish species reported to occur in the watershed, but not in the lake (as in Pearse1920) were not considered part of the lakes biota.