Area and mammalian elevational diversity

Elevational gradients hold enormous potential for understanding general properties of biodiversity. Like latitudinal gradients, the hypotheses for diversity patterns can be grouped into historical explanations, climatic drivers, and spatial hypotheses. The spatial hypotheses include the species-area effect and spatial constraint (mid-domain effect null models). I test these two spatial hypotheses using regional diversity patterns for mammals (non-volant small mammals and bats) along 34 elevational gradients spanning 24.4 degrees S-40.4 degrees N latitude. There was high variability in the fit to the species-area hypothesis and the mid-domain effect. Both hypotheses can be eliminated as primary drivers of elevational diversity. Area and spatial constraint both represent sources of error rather than mechanisms underlying these mammalian diversity patterns. Similar results are expected for other vertebrate taxa, plants, and invertebrates since they show comparable distributions of elevational diversity patterns to mammalian patterns.     

Reliability of a Higher-Taxon Approach to Richness, Rarity, and Composition Assessments at the Local Scale

Abstract:  A promising shortcut for quantifying species patterns is to use genera and families as surrogates of species. At large spatial scales, concurrence between patterns of richness, rarity, and composition of species and higher taxa is generally high. Only a few researchers, however, have examined this relationship at the local scale, which is frequently the relevant scale in land-use conflicts. We investigated the reliability of the higher-taxon approach in assessing patterns of species richness, rarity, and composition at the local scale. We studied diversity patterns of three commonly used surrogate taxa: vascular plants, ground-dwelling beetles, and moths. We conducted year-round field surveys for these taxa in the Jerusalem Mountains and the Judean foothills, Israel. Richness and composition of species were highly correlated with richness and composition of genera for all taxa. At the family level, correlations with richness and composition of species were much lower. Excluding monotypic genera and families did not affect these relations. Rarity representation based on higher taxa varied considerably depending on the taxon, and rarity scale and was weaker compared with richness and composition representation. Cumulative richness curves of species and genera showed similar patterns, leveling off at equivalent sampling efforts. Genus-level assessments were a reliable surrogate for local patterns of species richness, rarity, and composition, but family-level assessments performed poorly. The advantage of using coarse taxonomic scales in local diversity surveys is that it may decrease identification time and the need for experts, but it will not reduce sampling effort.     

Species Richness, Endemism, and the Choice of Areas for Conservation

Although large reserve networks will be integral components in successful biodiversity conservation, implementation of such systems is hindered by the confusion over the relative importance of endemism and species richness. There is evidence (  Prendergast et al. 1993) that regions with high richness for a taxon tend to be different from those with high endemism. I tested this finding using distribution and richness data for 368 species from Mammalia, Lasioglossum, Plusiinae, and Papilionidae. The study area, subdivided into 336 quadrats, was the continuous area of North America north of Mexico. I also tested the hypothesis that the study taxa exhibit similar diversity patterns in North America. I found that endemism and richness patterns within taxa were generally similar. Therefore, the controversy over the relative importance of endemism and species richness may not be necessary if an individual taxon were the target of conservation efforts. I also found, however, that richness and endemism patterns were not generally similar between taxa. Therefore, centering nature reserves around areas that are important for mammal diversity (the umbrella species concept) may lead to large gaps in the overall protection of biodiversity because the diversity and endemism of other taxa tend to be concentrated elsewhere. I investigated this further by selecting four regions in North America that might form the basis of a hypothetical reserve system for Carnivora. I analyzed the distribution of the invertebrate taxa relative to these regions and found that this preliminary carnivore reserve system did not provide significantly different protection for these invertebrates than randomly selected quadrats. I conclude that the use of Carnivora as an umbrella taxon is an unreliable method for invertebrate conservation.     

Global distribution of a key trophic guild contrasts with common latitudinal diversity patterns

Most hypotheses explaining the general gradient of higher diversity toward the equator are implicit or explicit about greater species packing in the tropics. However, global patterns of diversity within guilds, including trophic guilds (i.e., groups of organisms that use similar food resources), are poorly known. We explored global diversity patterns of a key trophic guild in stream ecosystems, the detritivore shredders. This was motivated by the fundamental ecological role of shredders as decomposers of leaf litter and by some records pointing to low shredder diversity and abundance in the tropics, which contrasts with diversity patterns of most major taxa for which broad-scale latitudinal patterns haven been examined. Given this evidence, we hypothesized that shredders are more abundant and diverse in temperate than in tropical streams, and that this pattern is related to the higher temperatures and lower availability of high-quality leaf litter in the tropics. Our comprehensive global survey (129 stream sites from 14 regions on six continents) corroborated the expected latitudinal pattern and showed that shredder distribution (abundance, diversity and assemblage composition) was explained by a combination of factors, including water temperature (some taxa were restricted to cool waters) and biogeography (some taxa were more diverse in particular biogeographic realms). In contrast to our hypothesis, shredder diversity was unrelated to leaf toughness, but it was inversely related to litter diversity. Our findings markedly contrast with global trends of diversity for most taxa, and with the general rule of higher consumer diversity at higher levels of resource diversity. Moreover, they highlight the emerging role of temperature in understanding global patterns of diversity, which is of great relevance in the face of projected global warming.     

Diversity and host range of foliar fungal endophytes: are tropical leaves biodiversity hotspots?

Fungal endophytes are found in asymptomatic photosynthetic tissues of all major lineages of land plants. The ubiquity of these cryptic symbionts is clear, but the scale of their diversity, host range, and geographic distributions are unknown. To explore the putative hyperdiversity of tropical leaf endophytes, we compared endophyte communities along a broad latitudinal gradient from the Canadian arctic to the lowland tropical forest of central Panama. Here, we use molecular sequence data from 1403 endophyte strains to show that endophytes increase in incidence, diversity, and host breadth from arctic to tropical sites. Endophyte communities from higher latitudes are characterized by relatively few species from many different classes of Ascomycota, whereas tropical endophyte assemblages are dominated by a small number of classes with a very large number of endophytic species. The most easily cultivated endophytes from tropical plants have wide host ranges, but communities are dominated by a large number of rare species whose host range is unclear. Even when only the most easily cultured species are considered, leaves of tropical trees represent hotspots of fungal species diversity, containing numerous species not yet recovered from other biomes. The challenge remains to recover and identify those elusive and rarely cultured taxa with narrower host ranges, and to elucidate the ecological roles of these little-known symbionts in tropical forests.     

Historical processes constrain patterns in global diatom diversity

There is a long-standing belief that microbial organisms have unlimited dispersal capabilities, are therefore ubiquitous, and show weak or absent latitudinal diversity gradients. In contrast, using a global freshwater diatom data set, we show that latitudinal gradients in local and regional genus richness are present and highly asymmetric between both hemispheres. Patterns in regional richness are explained by the degree of isolation of lake districts, while the number of locally coexisting diatom genera is highly constrained by the size of the regional diatom pool, habitat availability, and the connectivity between habitats within lake districts. At regional to global scales, historical factors explain significantly more of the observed geographic patterns in genus richness than do contemporary environmental conditions. Together, these results stress the importance of dispersal and migration in structuring diatom communities at regional to global scales. Our results are consistent with predictions from the theory of island biogeography and metacommunity concepts and likely underlie the strong provinciality and endemism observed in the relatively isolated diatom floras in the Southern Hemisphere.     

Large-scale biodiversity patterns in freshwater phytoplankton

Our planet shows striking gradients in the species richness of plants and animals, from high biodiversity in the tropics to low biodiversity in polar and high-mountain regions. Recently, similar patterns have been described for some groups of microorganisms, but the large-scale biogeographical distribution of freshwater phytoplankton diversity is still largely unknown. We examined the species diversity of freshwater phytoplankton sampled from 540 lakes and reservoirs distributed across the continental United States and found strong latitudinal, longitudinal, and altitudinal gradients in phytoplankton biodiversity, demonstrating that microorganisms can show substantial geographic variation in biodiversity. Detailed analysis using structural equation models indicated that these large-scale biodiversity gradients in freshwater phytoplankton diversity were mainly driven by local environmental factors, although there were residual direct effects of latitude, longitude, and altitude as well. Specifically, we found that phytoplankton species richness was an increasing saturating function of lake chlorophyll a concentration, increased with lake surface area and possibly increased with water temperature, resembling effects of productivity, habitat area, and temperature on diversity patterns commonly observed for macroorganisms. In turn, these local environmental factors varied along latitudinal, longitudinal, and altitudinal gradients. These results imply that changes in land use or climate that affect these local environmental factors are likely to have major impacts on large-scale biodiversity patterns of freshwater phytoplankton.     

A Mechanistic Approach to Evaluation of Umbrella Species as Conservation Surrogates

Abstract:  Although species with large area requirements are sometimes used as umbrella species, their general utility as conservation tools is uncertain. We surveyed the species diversity of birds, butterflies, carabids, and forest-floor plants in forest sites across an area (1600 km²) in which we delineated large breeding home ranges of Northern Goshawk ( Accipiter gentilis ). We tested whether protection of the home ranges could serve as an effective umbrella to protect sympatric species of the four taxa. We also used an empirical habitat model of occupancy of home range to examine mechanisms by which the Northern Goshawk acts as an umbrella species. Among species richness, abundance, and species composition of the four taxa, only abundance and species composition of birds differed between sites located inside and outside home ranges, which was due to greater abundance of bird species that were prey of Northern Goshawks inside the home ranges. Thus, although home range indicated areas with high abundance of certain bird prey species, it was not effective as an indicator of the species diversity of all four taxa. We also did not find any difference in species richness, abundance, and species composition between sites predicted as occupied and unoccupied using the habitat model. In contrast, when we selected sites on the basis of each habitat variable in the model, habitat variables that selected sites either in agricultural or forested landscapes encompassed sites with high species richness or particular species composition. This result suggests that the low performance of the Northern Goshawk as an umbrella species is due to this species' preference for habitat in both agricultural and forested landscapes. Species that can adjust to changes in habitat conditions may not act as effective umbrella species despite having large home ranges.     

Trends over time in tree and seedling phylogenetic diversity indicate regional differences in forest biodiversity change

Changing climate conditions may impact the short-term ability of forest tree species to regenerate in many locations. In the longer term, tree species may be unable to persist in some locations while they become established in new places. Over both time frames, forest tree biodiversity may change in unexpected ways. Using repeated inventory measurements five years apart from more than 7000 forested plots in the eastern United States, we tested three hypotheses: phylogenetic diversity is substantially different from species richness as a measure of biodiversity; forest communities have undergone recent changes in phylogenetic diversity that differ by size class, region, and seed dispersal strategy; and these patterns are consistent with expected early effects of climate change. Specifically, the magnitude of diversity change across broad regions should be greater among seedlings than in trees, should be associated with latitude and elevation, and should be greater among species with high dispersal capacity. Our analyses demonstrated that phylogenetic diversity and species richness are decoupled at small and medium scales and are imperfectly associated at large scales. This suggests that it is appropriate to apply indicators of biodiversity change based on phylogenetic diversity, which account for evolutionary relationships among species and may better represent community functional diversity. Our results also detected broadscale patterns of forest biodiversity change that are consistent with expected early effects of climate change. First, the statistically significant increase over time in seedling diversity in the South suggests that conditions there have become more favorable for the reproduction and dispersal of a wider variety of species, whereas the significant decrease in northern seedling diversity indicates that northern conditions have become less favorable. Second, we found weak correlations between seedling diversity change and latitude in both zones, with stronger relationships apparent in some ecoregions. Finally, we detected broadscale seedling diversity increases among species with longer-distance dispersal capacity, even in the northern zone, where overall seedling diversity declined. The statistical power and geographic extent of such analyses will increase as data become available over larger areas and as plot measurements are repeated at regular intervals over a longer period of time.     

Poleward shifts in winter ranges of North American birds

Climate change is thought to promote the poleward movement of geographic ranges; however, the spatial dynamics, mechanisms, and regional anthropogenic drivers associated with these trends have not been fully explored. We estimated changes in latitude of northern range boundaries, center of occurrence, and center of abundance for 254 species of winter avifauna in North America from 1975 to 2004. After accounting for the effect of range size and the location of the northern boundary, positive latitudinal trends were evident for the northern boundary (1.48 km/yr), center of occurrence (0.45 km/yr), and center of abundance (1.03 km/yr). The northern boundary, when examined across individual species, had the most variable trends (SD = 7.46 km/yr) relative to the center of occurrence (SD = 2.36 km/yr) and center of abundance (SD = 5.57 km/yr). Trends did not differ based on migratory status, but there was evidence that trends differed for species with ranges centered in the southern vs. northern portion of the study area. Species occurred more sporadically over time at northern range boundaries, and northern boundaries were associated with a concentration of colonization and extirpation events, with a greater prevalence of colonization events likely promoting poleward trends. Regional anthropogenic drivers explained approximately 8% of the trend for the northern boundary, 14% for the center of occurrence, and 18% for the center of abundance; however, these effects were localized in the northern portion of species' ranges and were associated with distributional changes within ranges, primarily abundance, producing patterns that mimicked poleward movements. We conclude that poleward distributional shifts represent the interaction between climate change and regional factors whose outcome is determined by the scale of the analysis and the biotic and abiotic features in the region, and how anthropogenic activities have impacted these features.     

Beyond Species Richness: Community Similarity as a Measure of Cross-Taxon Congruence for Coarse-Filter Conservation

Abstract:  The use of a surrogate taxon in conservation planning has become questionable because recent evidence suggests that the correlation of species richness between pairs of taxa is highly variable both taxonomically and geographically. Species richness is only one measure of species diversity, however, and recent studies suggest that investigations of cross-taxon congruence should consider a broader range of assessment techniques. The cross-taxon congruence of community similarity between sites among taxa has rarely been examined and may be the most relevant measure of species diversity in the context of coarse-filter conservation strategies. We examined cross-taxon congruence patterns of species richness and community similarity (Bray-Curtis similarity) among birds, butterflies, and vascular plants in montane meadow habitats in the Greater Yellowstone Ecosystem. Although patterns of species richness (Spearman rank correlation) varied between taxa, we consistently found a positive correlation in community similarity (Mantel test) between all pair-wise comparisons of the three taxa (e.g., sites with similar bird communities also had similar butterfly communities). We suggest that the success of a surrogate taxon depends on the technique used to assess surrogacy and the specific approach to conservation planning. In the context of coarse-filter conservation, measures of community similarity may be more appropriate than measures of species richness. Furthermore, the cross-taxon congruency of community similarity in our study suggests that coarse-filter conservation may be tenable in montane meadow communities.     

Trade-Offs between Species Conservation and the Size of Marine Protected Areas

Abstract:  Moving from single-species- to ecosystem-based management requires an understanding of how community-level attributes such as diversity change with area. We used survey data from bottom trawls to examine spatial patterns of species richness in U.S. Pacific coastal fishes. Specifically, we generated and compared species–area relationships (SARs) for species classified into several groups on the basis of maximum body size, trophic level, diet, maximum depth, geographic affinity, and taxonomic order. Because SARs among groups were not parallel and z values varied significantly for several groups, groups of species were under- or overrepresented (depending on the size of the area) relative to their proportions in the entire community (i.e., entire U.S. Pacific coast). In this way, differences in SARs help demonstrate trade-offs between species representation and coastal area and suggest strategies (such as targeting the protection of habitats and locations where a particular species or groups of species are maximized) that may minimize the size of marine protected areas (MPAs) but protect diversity at the level of the community and functional group.     

Geographical gradients of marine herbivorous fishes: patterns and processes

We present new data and the first rigorous analysis of latitudinal and thermal gradients of diversity, density and biomass of marine herbivorous fishes and review proposed explanatory mechanisms. Consistently, negative relationships between latitude, and positive relationships between sea surface temperature (SST), and relative richness and relative abundance of herbivorous fishes were found worldwide. Significant differences in the strength of gradients of richness and abundance with latitude and SST between tropical and extratropical zones were found consistently across ocean basins. Standardized sampling along the western Atlantic also showed negative relationships between latitude and total density and biomass. The trends, however, are driven by different components of the fish assemblages (i.e., scarids in the Caribbean and acanthurids in Brazil). Patterns of abundance along thermal gradients, generally associated with extensive latitudinal gradients, also were found at the local scale. Feeding rate of the ocean surgeonfish Acanthurus bahianus decreases with temperature more rapidly than the mean metabolic rate of teleost fishes. This relationship suggests a temperature-related physiological constraint. From the new standardized and comparative data presented and the review of the explanatory hypotheses, we conclude that temperature-related feeding and digestive processes are most likely involved in the distribution patterns of herbivorous fishes. Electronic Supplementary Material Supplementary material is available for this article at     

Modelling abiotic indicators when obtaining spatial predictions of species richness

When conservation biologists formulate strategy used in decisions concerning the locations of new national parks, refuges and reserves, accurate information about species richness and spatialpatterns of species distributions can be critical. Recent research has demonstrated that spatial models and bioindicator taxa can be quite useful for determining generalized spatial patterns of unrelated taxa on a continental scale. In this research, I incorporate abiotic effects, in this case altitudinal relief, into both the mean and the covariance structures of the spatial prediction model. I use bird species data collected in the Indian subcontinent and cross-validation techniques to illustrate the degree of improvement in prediction accuracy engendered by using theabiotic factor and the modified spatial models.     

Habitat biodiversity as a determinant of fish community structure on coral reefs

Increased habitat diversity is often predicted to promote the diversity of animal communities because a greater variety of habitats increases the opportunities for species to specialize on different resources and coexist. Although positive correlations between the diversities of habitat and associated animals are often observed, the underlying mechanisms are only now starting to emerge, and none have been tested specifically in the marine environment. Scleractinian corals constitute the primary habitat-forming organisms on coral reefs and, as such, play an important role in structuring associated reef fish communities. Using the same field experimental design in two geographic localities differing in regional fish species composition, we tested the effects of coral species richness and composition on the diversity, abundance, and structure of the local fish community. Richness of coral species overall had a positive effect on fish species richness but had no effect on total fish abundance or evenness. At both localities, certain individual coral species supported similar levels of fish diversity and abundance as the high coral richness treatments, suggesting that particular coral species are disproportionately important in promoting high local fish diversity. Furthermore, in both localities, different microhabitats (coral species) supported very different fish communities, indicating that most reef fish species distinguish habitat at the level of coral species. Fish communities colonizing treatments of higher coral species richness represented a combination of those inhabiting the constituent coral species. These findings suggest that mechanisms underlying habitat-animal interaction in the terrestrial environment also apply to marine systems and highlight the importance of coral diversity to local fish diversity. The loss of particular key coral species is likely to have a disproportionate impact on the biodiversity of associated fish communities.     

High genetic diversities and complex genetic structure in an Indo-Pacific tropical reef fish (Chlorurus sordidus): evidence of an unstable evolutionary past?

Historical sea level fluctuations have influenced the genetic structure and evolutionary history of marine species and examining widespread species across their species ranges may elucidate some of these effects. Chlorurus sordidus is a common and widespread parrotfish found on coral reefs throughout the Indo-central Pacific. We used phylogenetic, phylogeographic, and cladistic analyses to examine the genetic composition and population structure of this species across most of its latitudinal range limits. We sequenced 354 bp of the mitochondrial control region I in 185 individuals from nine populations. Populations of C. sordidus displayed high levels of genetic diversity, similar to those recorded for widespread pelagic fish species, but much greater nucleotide diversity values than those previously recorded for other demersal reef fishes. Both phylogenetic and phylogeographic analyses detected strong genetic subdivision at the largest spatial scale (i.e. among oceans). The Pacific Ocean was characterised by weak population genetic structure. Separation of the Hawaiian location from other Pacific and West Indian Ocean sites was evident in phylogenetic analyses, but not from analysis of molecular variance. NCA and isolation-by-distance tests suggested that the genetic structure of this species was the result of multiple contemporary and historical processes, including long-distance colonisation and range expansion arising from fluctuating sea levels, limited current gene flow, and isolation by distance. This pattern is to be expected when historically fragmented populations come into secondary contact. We suggest the patterns of population genetic structure recorded in C. sordidus are caused by large local population sizes, high gene flow, and a recent history of repeated fragmentation and remixing of populations resulting from fluctuating sea levels.Communicated by M.S. Johnson, Crawley     

Benefits to poorly studied taxa of conservation of bird and mammal diversity on islands

Protected area delineation and conservation action are urgently needed on marine islands, but the potential biodiversity benefits of these activities can be difficult to assess due to lack of species diversity information for lesser known taxa. We used linear mixed effects modeling and simple spatial analyses to investigate whether conservation activities based on the diversity of well‐known insular taxa (birds and mammals) are likely to also capture the diversity of lesser known taxa (reptiles, amphibians, vascular land plants, ants, land snails, butterflies, and tenebrionid beetles). We assembled total, threatened, and endemic diversity data for both well‐known and lesser known taxa and combined these with physical island biogeography characteristics for 1190 islands from 109 archipelagos. Among physical island biogeography factors, island area was the best indicator of diversity of both well‐known and little‐known taxa. Among taxonomic factors, total mammal species richness was the best indicator of total diversity of lesser known taxa, and the combination of threatened mammal and threatened bird diversity was the best indicator of lesser known endemic richness. The results of other intertaxon diversity comparisons were highly variable, however. Based on our results, we suggest that protecting islands above a certain minimum threshold area may be the most efficient use of conservation resources. For example, using our island database, if the threshold were set at 10 km² and the smallest 10% of islands greater than this threshold were protected, 119 islands would be protected. The islands would range in size from 10 to 29 km² and would include 268 lesser known species endemic to a single island, along with 11 bird and mammal species endemic to a single island. Our results suggest that for islands of equivalent size, prioritization based on total or threatened bird and mammal diversity may also capture opportunities to protect lesser known species endemic to islands. Beneficios de los Taxa Poco Estudiados para la Conservación de la Diversidad de Aves y Mamíferos en Islas     

A comparative analysis of the photobiology of Zooxanthellae and Zoochlorellae symbiotic with the temperate clonal anemone Anthopleura elegantissima (Brandt) I. Effect of temperature

Throughout its geographic range, the temperate-zone anemone Anthopleura elegantissima is the host of one or both of two distinctively different symbiotic microalgae: a dinoflagellate Symbiodinium (zooxanthellae, ZX) and a chlorophyte (zoochlorellae, ZC). Given the broad vertical intertidal and latitudinal range of this anemone, we investigated the role of temperature in determining whether A. elegantissima supports one algal symbiont over the other and whether temperature regulates the observed distributions of natural populations of ZX and ZC. Temperature appears to be a key factor in regulating both the photophysiology and metabolism of this algal–cnidarian association. In anemones containing ZX, neither algal densities nor chlorophyll content varied with temperature (6–24 °C); in contrast, anemones with ZC displayed reduced densities and chlorophyll content at the highest temperature treatment (24 °C). Both ZX and ZC photosynthetic rates were directly related to temperature, as were anemone respiration rates. The higher photosynthetic rates, maintenance of a stable algal density and chlorophyll content, and higher potential contribution of algal carbon toward animal respiration (CZAR) suggest that the ZX are the more viable symbiont as temperature increases, but we suggest alternative reasons why ZC are preserved in this symbiotic association. Elevated temperatures reduce ZC densities and chlorophyll, suggesting that higher temperatures affect this relationship in a negative fashion, presumably due to a higher cost of maintaining ZC by the association; alternatively, these costs may be affiliated with the deterioration of the ZC themselves. These results suggest that temperature may be one of the most significant environmental parameters that sets the intertidal microhabitat and latitudinal distribution patterns of the two algal taxa observed in the field. Received: 2 November 1998 / Accepted: 25 October 2000     

Recovery of Faunal Communities During Tropical Forest Regeneration

Abstract:  As mature tropical forests are cleared, secondary forests may play an important role in the conservation of animal species, depending on how fast animal communities recover during forest regeneration. I reviewed published studies on the recovery of animal species richness and composition during tropical forest regeneration. In 38 of the 39 data sets I examined, conversion of forest to agriculture or pasture substantially reduced species richness. Given suitable conditions for forest recovery, the species richness of the animal taxa considered can be predicted to resemble that of mature forests roughly 20–40 years after land abandonment. At least for ants and birds, however, recovery of species composition appears to take substantially longer than recovery of species richness. Because species richness for many taxa appears to recover relatively rapidly in secondary forests, conservation of secondary forests may be an effective investment in future diversity. The slower recovery of species composition indicates, however, that some species will require stands of mature forest to persist.     

Physical and biological disturbances interact differently with productivity: effects on floral and faunal richness

Physical and biological disturbances are ecological processes affecting patterns in biodiversity at a range of scales in a variety of terrestrial and aquatic systems. Theoretical and empirical evidence suggest that effects of disturbance on diversity differ qualitatively and quantitatively, depending on levels of productivity (e.g., the dynamic equilibrium model). In this study we contrasted the interactive effects between physical disturbance and productivity to those between biological disturbance and productivity. Furthermore, to evaluate how these effects varied among different components of marine hard-substratum assemblages, analyses were done separately on algal and invertebrate richness, as well as richness of the whole assemblage. Physical disturbance (wave action) was simulated at five distinct frequencies, while biological disturbance (grazing periwinkles) was manipulated as present or absent, and productivity was manipulated as high or ambient. Uni- and multivariate analyses both showed significant effects of physical disturbance and interactive effects between biological disturbance and productivity on the composition of assemblages and total species richness. Algal richness was significantly affected by productivity and biological disturbance, whereas invertebrate richness was affected by physical disturbance only. Thus, we show, for the first time, that biological disturbance and physical disturbance interact differently with productivity, because these two types of disturbances affect different components of assemblages. These patterns might be explained by differences in the distribution (i.e., press vs. pulse) and degree of selectivity between disturbances. Because different types of disturbance can affect different components of assemblages, general ecological models will benefit from using natural diverse communities, and studies concerned with particular subsets of assemblages may be misleading. In conclusion, this study shows that the outcome of experiments on effects of disturbance and productivity on diversity is greatly influenced by the composition of the assemblage under study, as well as on the type of disturbance that is used as an experimental treatment.