Facilitation of periphyton production by tadpole grazing: functional differences between species

1. This study examined how interactions between resources that vary in edibility, and herbivores that vary in ability to acquire resources, control primary productivity. In a northern California river, grazing on Cladophora glomerata , a relatively inedible filamentous green alga, and its more nutritious epiphytic diatoms, was manipulated by exposing cobbles to tadpoles ( Rana boylii or Hyla regilla ) or excluding tadpoles.
2. Rana indirectly facilitated Cladophora by removing diatoms, whereas Hyla did not significantly change biomass relative to controls. Algal ash-free dry mass on cobbles in Rana treatments was 65 and 72% greater than on controls in two years of investigation (1991 and 1993). Rana decreased epiphytic diatom biovolume by 56% and detritus by 87%.
3. Because nitrogen excretion rates of Hyla and Rana were similar, the differences in effect between the two species were probably due to their roles as consumers rather than as recyclers.
4. The net effect of Rana on periphyton was a 10% increase in areal specific primary productivity (mg O₂ h^–1 m^–2); Hyla caused an 18% decrease. Rana decreased biomass-specific productivity (mg O₂ h^–1 g^–1) 44%; Hyla had no effect.
5. In tadpole exclosures, grazers such as baetid mayfly larvae (mostly Centroptilum sp.) were 4.7 (1991) and 1.8 (1993) times more abundant, and midge larvae (Chironomidae) were 2.5 (1991) and 2 (1993) times more abundant than in Rana enclosures. Invertebrate assemblages in Hyla enclosures, however, were similar to exclosures. Few predatory insects and fish colonized Rana enclosures. Path analyses indicated that Rana affected macroinvertebrates via both interference and exploitation of epiphytic diatoms.     

Interactions between neighboring algae and snail grazing in structuring microdistribution patterns of periphyton

The micro-distribution of periphyton (filamentous algae) on homogeneous substrates was examined in experimental tanks with and without the pressure of grazing snails. The growth of periphyton attached to artificial substrate was estimated at a small spatial scale (9.3 mm×9.3 mm cells) by varying the number of grazers (0, 5, or 10 snails per tank), using image processing analysis without removing the periphyton. The results suggest that periphyton growth within a cell was negatively affected by the biomass of periphyton in the cell but was positively affected by the biomass of periphyton in neighboring cells. A semivariogram analysis indicated that spatial heterogeneity increased with increasing grazing pressure. The size of patches was not clearly related to the number of snails, but there was a tendency for relative patch size to increase with snail density. Computer simulations were also conducted to examine factors affecting the degree of spatial heterogeneity. The simulation studies indicated that snails should graze a site that was previously grazed in order to produce the observed spatial heterogeneity of periphyton. The results also indicated that the positive effects of neighboring periphyton on the growth of algae might create patches. The interactions among neighboring algae and snail grazing might be an important factor creating the spatial heterogeneity of periphyton even on homogeneous substrates.     

Alterations in the biomass-specific productivity of periphyton assemblages mediated by fish grazing

1. In an experimental flume, we examined the effects of a biomass reduction and alteration of taxonomic composition, because of grazing by the fish Plecoglossus altivelis, on the net biomass accumulation of periphyton. 2. Grazed and ungrazed assemblages with different biomass and taxonomic composition were first prepared in fish enclosures and exclosures, respectively. These assemblages were then set out in the flume and incubated for 2 days under grazing‐free conditions to examine (i) the relationship between biomass and biomass accumulation rate and (ii) the effect of taxonomic composition on the relationship between these two. 3. The grazed and ungrazed assemblages were dominated by upright filamentous cyanobacteria and diatoms, respectively. The rate of biomass accumulation decreased with increasing periphyton biomass in both the grazed and ungrazed assemblages, and was lower in the grazed than the ungrazed assemblages at any biomass level. 4. The results showed that the reduction in biomass and the alteration of taxonomic composition due to fish grazing have opposite effects on biomass‐specific productivity. Biomass accumulation rate increased in response to biomass reduction, although a shift in dominance from diatoms to upright filamentous cyanobacteria decreased the overall productivity of the periphyton.     

Facilitation of clear-water conditions in shallow lakes by macrophytes: differences between charophyte and angiosperm dominance

A number of mechanisms result in a feedback between water clarity and macrophytes and, consequently, the occurrence of alternative stable states in shallow lakes. We hypothesize that bottom-up mechanisms and interactions within the benthic food web are more important in a charophyte-dominated clear-water state, while top-down mechanism and interactions in the planktonic food web prevail at angiosperm dominance. Charophytes, which dominate at lower nutrient concentrations and develop higher densities than most angiosperms, can have a higher influence on sedimentation, resuspension, and water column nutrients. During dominance of dense submerged vegetation like charophytes, zooplankton can be hampered by low food quality and quantity and by high predation pressure from juvenile fish, which in turn are favoured by the high refuge potential of this vegetation. Grazing pressure from zooplankton on phytoplankton can therefore be low in charophytes, but the main feedback in angiosperm-dominated ecosystems. Charophytes offer a higher surface than most angiosperms to periphyton, which favors benthic invertebrates. These support macrophytes by grazing periphyton and constitute a central link in a trophic cascade from fish to periphyton and macrophytes. To test these hypotheses, more experiments and field measurements comparing the effect of charophytes and angiosperms on water clarity are needed.     

Trophic linkages between periphyton and grazing macroinvertebrates in rivers with different levels of catchment development

Stable isotope signatures (δ¹³C and δ¹⁵N) were used to compare trophic linkages between epilithic periphyton and three families of macroinvertebrates (Baetidae, Leptophlebiidae and Gripopterygidae) in riffles of two rivers with developed catchments (including agriculture, urbanization, impoundment and flow regulation) and two with undeveloped catchments (native forest with no major impoundments) in the Murrumbidgee River system, New South Wales, Australia. Periphyton had much higher average δ¹⁵N values and lower average C:N ratios in the developed rivers than in the undeveloped rivers, probably because of the combined effects of nutrient enrichment, upstream impoundment and alteration of riparian vegetation. The invertebrates were generally slightly depleted in ¹³C and ¹⁵N relative to expected values if they were assimilating whole periphyton alone, which suggests that they were assimilating periphyton components selectively or also consuming other foods. The match in isotope signatures between periphyton and invertebrates was only slightly weaker in the developed than in the undeveloped rivers, suggesting that development did not greatly disrupt trophic linkages between periphyton and these invertebrates. Handling editor: M. Power     

Effects of tadpole grazing on periphytic algae in ponds

Tadpole impact on periphytic algae was estimated with an exclosure experiment in the field. Algae growth on control (exposed) and experimental (netted-in) Perspex plates was measured in 12 ponds in 1991 and 8 in 1993.An index of snail (Lymnea and Planorbis)presence in the ponds was used to correct their effect. Grazing pressure (difference in amount of algae between control and experimental plates)differed between ponds. The difference was correlated to amount of Rana tadpoles present in the pond. At the higher densities of tadpoles observed in the study ponds, the algae standing crop on exposed plates was about 50–75% of that on netted-in plates. These result demonstrate that there is potential for effects of exploitation competition in ponds with a high tadpole density. This revised version was published online in July 2006 with corrections to the Cover Date.     

The effect of grazing and nutrient supply on periphyton associated bacteria

The effects of nutrient additions and grazing by macro-invertebrates on periphyton-associated algae and bacteria were studied by performing an enclosure experiment on three occasions from early spring to summer at mesotrophic Lake Erken and V?dd?, at the Swedish Baltic coast. There were significant interactions between nutrient additions and grazing on bacterial biomass and specific activity in Lake Erken. Thus, the importance of either bottom-up or top-down effects could not be singled out. Bacterial biomass increased with enrichment only in the absence of grazers. Grazer presence tended to increase bacterial biomass in ambient nutrient conditions, but to decrease bacterial biomass under enrichment. For specific activity the positive response to enrichment was restricted to grazer presence. Hence, grazing by macro-invertebrates may have an indirect positive effect on bacterial activity by enhancing nutrient conditions through their feeding activities and/or fecal pellets production. In addition, we found a significant relationship between bacterial production and chlorophyll a at both sites. This relationship weakened in the presence of macro-invertebrates. Thus, the importance of internal nutrient regeneration by bacteria and algae decreased, possibly due to increased nutrient availability, in the presence of macro-invertebrate grazers.     

Control of plant growth by nitrogen: differences between cereals and broadleaf species

Abstract. In four dicotyledonous species low levels of N strongly inhibited leaf expansion during the day but had little or no effect at night. In contrast, daytime and night-time expansion were equally affected in four cereal species. The results support the general concept that in dicotyledons, N controls leaf expansion through its effects on hydraulic conductivity. In such N-limited plants, water deficits generated by transpiration may inhibit daytime cell expansion. In cereals, cell expansion and transpiration occur in separate zones of the leaf and are apparently unrelated.
Growth analysis showed that low levels of N inhibited leaf area growth more strongly in dicotyledons than in cereals, but had similar effects on net assimilation rates of plants in the two groups. As a result, dry matter production was more efficient in cereals than in dicotyledons when N was limiting.     

The influence of periphyton biomass and density on grazing in Physella virgata

We studied how differences in periphyton colonization interval and snail density affected grazing rates in Physella virgata, and whether snails controlled periphyton biomass. Both egestion rates and incorporation rates of ¹⁴C labeled periphyton were estimated in laboratory experiments. Periphyton biomass increased with field colonization interval in all experiments, but did not consistently influence estimates of grazing rate. However, increased periphyton abundance in one of the experiments could still explain higher grazer rates in that year, although larger snail body size is a confounding explanation. Increased snail density also resulted in decreased grazing rates, as observed in earlier studies with this snail species, as well as in studies with other snail grazers. Our results suggest grazing rates and resulting impacts may change seasonally with variation in either periphyton biomass, grazer life-history stage or population density.     

Body shape and differences between species

The variation of body shape among prosimians is reviewed. Special emphasis is placed on the selective advantages, that is the mechanical reasons, to which variants of the locomotor apparatus can be traced back. There are differences found in the cheiridia, but at present they cannot be explained in terms of mechanics; there is nearly no knowledge about the mechanical meaning of their diversity. Myological characteristics of taxa can be explained mechanically, but this has not yet been done. Well known are variations of body proportions. These discriminate higher taxa, and are largely coincident with the often-used locomotor categories. In spite of this, there are only few sound arguments about the real biomechanic value of characteristic proportions for a given locomotor mode. What is known on this field, is reviewed. Progress can be made only, if the mechanical conditions, set by postural behavior and locomotion, are understood completely. The subtle distinctions between lower taxonomic units can normally be identified only on the basis of detailed and quantified analyses of movements on one hand, and of biometrics on the other. In the few cases in which such studies have been made, the differences of morphology fit to the mechanical requirements of locomotion which also differ only in quantitative details.     

Immunoelectrophorecal differences between phytopathogenic Fusarium species

Immunoelectrograms of certain tested Fusarium spp revealed that each has a characteristic pattern and showed the relatedness between F. oxysporum on the one hand and F. culmorum and F. semitectum on the other. The B₄ protein fraction is common to the former pair, while α_1a protein fraction links the second.     

Variations in species and functional plant diversity along climatic and grazing gradients

Different components of biodiversity may vary independently of each other along environmental gradients giving insights into the mechanisms that regulate species coexistence. In particular, the functional diversity (FD) or the presence of rare or endemic species in natural assemblages do not necessarily increase with species diversity. We studied if different components of plant species diversity (species richness, Simpson diversity, evenness) varied similarly to FD (measured as a generalization of the Simpson index) and rarity along grazing intensity and climatic gradients. Plots under different sheep grazing regimes (high and low intensity, abandonment) were surveyed in five locations along a climatic gradient in north-eastern Spain, from semi-arid lowland to moist upland locations. Variation in species diversity, functional diversity and rarity followed different patterns. Species diversity was lowest in water-stressed environments (arid locations and southern aspects) and increased with grazing more makedly in humid locations. The FD was comparable between the most species-poor and species-rich locations and decreased with grazing in the moistest location, i.e. where species diversity markedly increased. The FD did not show a strong correlation with species richness nor with the Simpson index and less specious communities could show the highest functional diversity. The rarest species in the region were more frequently found in the abandoned areas, which held the lowest species diversity. Consequently, the mechanisms that enhance the diversity of species do not necessarily support a functional differentiation among those species or the maintenance of rare species in a region. We hypothesize that the degree of dependence of functional diversity on species diversity might be mostly related to the amplitude of the species' traits pool and on how species partition the niche space available.     

Nutrient loading and grazing by the minnow Phoxinus erythrogaster shift periphyton abundance and stoichiometry in mesocosms

1. Anthropogenic activities in prairie streams are increasing nutrient inputs and altering stream communities. Understanding the role of large consumers such as fish in regulating periphyton structure and nutritional content is necessary to predict how changing diversity will interact with nutrient enrichment to regulate stream nutrient processing and retention. 2. We characterised the importance of grazing fish on stream nutrient storage and cycling following a simulated flood under different nutrient regimes by crossing six nutrient concentrations with six densities of a grazing minnow (southern redbelly dace, Phoxinus erythrogaster) in large outdoor mesocosms. We measured the biomass and stoichiometry of overstory and understory periphyton layers, the stoichiometry of fish tissue and excretion, and compared fish diet composition with available algal assemblages in pools and riffles to evaluate whether fish were selectively foraging within or among habitats. 3. Model selection indicated nutrient loading and fish density were important to algal composition and periphyton carbon (C): nitrogen (N). Nutrient loading increased algal biomass, favoured diatom growth over green algae and decreased periphyton C : N. Increasing grazer density did not affect biomass and reduced the C : N of overstory, but not understory periphyton. Algal composition of dace diet was correlated with available algae, but there were proportionately more diatoms present in dace guts. We found no correlation between fish egestion/excretion nutrient ratios and nutrient loading or fish density despite varying N content of periphyton. 4. Large grazers and nutrient availability can have a spatially distinct influence at a microhabitat scale on the nutrient status of primary producers in streams.     

Cyclops predation on ciliates: species.specific differences and functional responses

Experiments were conducted to measure to what extent cyclopoidcopepods ingest ciliated protists. Five freshwater ciliate species,ranging in size from 22 to 120 µm diameter, were testedwith two species of cyclopoids: Cyclops abyssorum and Cyclopskolensis. Ingestion rates were measured by radiolabeling ciliateswith ¹⁴C, and from these, functional response curves (the changein ingestion rate with changing cell densities) were constructed.Cyclopoids ingest ciliates with very high estimated maximalrates of >200 cells cyclopoid^–1 h^–1 However,there are large differences in ingestion rates that are notpredictable by the size of predator or prey. One ciliate speciesof intermediate size, Coleps hirtus, is nearly immune from cyclopoidpredation at all measured ciliate densities. Three other smallciliate species that move in rapid jumps elicit Honing type3 functional responses, with very little change in ingestionrates at low ciliate densities. Thus, while cyclopoids are capableof having a very considerable impact on ciliate populations,some ciliate species appear to have behavioral, morphologicalor chemical defenses to reduce their vulnerability. This callsinto question the practice of considering ciliates a homogeneousgroup when constructing food web models.     

Unrecognized coral species diversity masks differences in functional ecology

Porites corals are foundation species on Pacific reefs but a confused taxonomy hinders understanding of their ecosystem function and responses to climate change. Here, we show that what has been considered a single species in the eastern tropical Pacific, Porites lobata, includes a morphologically similar yet ecologically distinct species, Porites evermanni. While P. lobata reproduces mainly sexually, P. evermanni dominates in areas where triggerfish prey on bioeroding mussels living within the coral skeleton, thereby generating asexual coral fragments. These fragments proliferate in marginal habitat not colonized by P. lobata. The two Porites species also show a differential bleaching response despite hosting the same dominant symbiont subclade. Thus, hidden diversity within these reef-builders has until now obscured differences in trophic interactions, reproductive dynamics and bleaching susceptibility, indicative of differential responses when confronted with future climate change.     

Determination of the periphyton production by means of the glass slide method

Summary By means of the glass slide method a study of the periphyton production was performed in the Sedlice Reservoir (Bohemia, Czechoslovakia) during the period July 30, 1958 – June 8th, 1959.The gravimetric mass analysis of the periphyton rendered at 9 sampling dates in 4–6 weeks intervals results for 4 depths (1, 3, 6, 9 m) including the zone of producers, the transitory zone, zone of consumers as well as the benthic zone.The air-dry weight and the dry weight (105°C) showed almost identical results. The dry weight formed in the average 8,0 % of the wet weight. The following values were calculated on the basis of the literature data: the ash-free dry weight by subtracting 20 % from the dry weight, the N-content as 7 % of the ash-free dry weight and the chlorophyll a content as 1,3 % of the ash-free dry weight.The biomass of the periphyton ranged between 0 and 491 mg/sq. dm of the ash-free dry weight with an average value 127,0. The highest values occurred in the depth of 1 m, the lowest near the bottom.The net production rate ranged between 10 and 1010 mg/sq.m/day of the ash-free dry weight with an average value 213. The highest mean value occurred in the depth of 6 m (280), the lowest in the depth of 9 m (91 mg/sq.m/day). The greatest production rates were found in March at all depths.The population turnover was different in each depth. In the depth of 1 m it equals 69,6 days, in 3 m 53,6 days, in 6 m 45,2 days, in 9 m 90,9 days. As a mean 59,6 = 60 days can be accepted. These are the longest turnover values determined at anyelse periphyton community. In the Sedlice Reservoir the periphyton renewes itself 6 times in the course of the year.All results gained were compared with similar data published by different authors. An attempt of correlation of these production data was undertaken, but no clear and reliable relationships could be detected.

---

Zusammenfassung Unter Anwendung der Glassplattenmethode wurde eine Untersuchung der Produktion des Aufwuchses in der Sedlice-Talsperre (Böhmen, Tschechoslowakei) in der Zeit vom 30. Juli 1958 bis 8. Juni 1959 durchgeführt.Die angewendete gravimetrische Globalmethode brachte an 9 Entnahmetagen (in 4–6 Wochen-Intervalen) Ergebnisse für 4 Tiefen (1, 3, 6, 9 m), die die Zone der Produzenten, die Übergangszone, die Zone der Konsumenten sowie die benthische Zone darstellen.Das Lufttrockengewicht und das Trockengewicht bei 105°C ergaben fast identische Ergebnisse. Das Trockengewicht stellte im Durchschnitt 8,0 % des Feuchtgewichtes dar. Die folgenden Werte wurden auf Grund von Literaturangaben ausgerechnet: das aschenfreie Trockengewicht durch die Subtraktion von 20 % vom Trockengewicht, der Stickstoffgehalt als 7 % des aschenfreien Trockengewichtes und der Chlorophyllgehalt als 1,3 % des aschenfreien Trockengewichtes.Die Biomasse des Aufwuchses schwankte zwischen 0 und 491 mg/m² des aschenfreien Trockengewichtes und zeigte den durchschnittlichen Wert 127,0. Die höchsten Werte kamen in der Tiefe 1 m und die niedrigsten in 9 m vor.Die Netto-Produktion des Aufwuchses schwankte zwischen 10 und 1010 mg/m²/Tag aschenfreier Trockensubstanz und zeigte den durchschnittlichen Wert 213,0. Die höchsten Durchschnittswerte waren in der Tiefe 6 m (280) und die niedrigsten in 9 m (91 mg/m²/Tag) zu verzeichnen. Im März fand sich in allen Tiefen die höchste Produktion.Der Erneuerungskoeffizient änderte sich mit der Tiefe. In 1 m betrug er 69,6 Tage, in 3 m 53,6 Tage, in 6 m 45,2 Tage, in 9 m 90,9 Tage und im Durchschnitt 59,6 = 60 Tage. Diese Werte sind als die längsten, die irgendwann im Aufwuchse ermittelt wurden, anzusehen. Das heisst, dass sich der Aufwuchs in der Sedlice-Talsperre nur sechsmal im Jahre erneuert.Alle erzielten Resultate wurden mit ähnlichen Literaturangaben verschiedener Autoren verglichen. Es wurde ein Versuch unternommen, alle verfügbaren Produktionswerte zur Korrelation zu bringen, eine klare Abhängigkeit konnte jedoch nicht festgestellt werden.

     

Species delimitation in systematics: inferring diagnostic differences between species

Species are fundamental units in studies of systematics, biodiversity and ecology, but their delimitation has been relatively neglected methodologically. Species are typically circumscribed based on the presence of fixed (intraspecifically invariant or non-overlapping) diagnostic morphological characters which distinguish them from other species. In this paper, we argue that determining whether diagnostic characters are truly fixed with certainty is generally impossible with finite sample sizes and we show that sample sizes of hundreds or thousands of individuals may be necessary to have a reasonable probability of detecting polymorphisms in diagnostic characters at frequencies approaching zero. Instead, we suggest that using a non-zero frequency cut-off may be a more realistic and practical criterion for character-based species delimitation (for example, allowing polymorphisms in the diagnostic characters at frequencies of 5% or less). Given this argument, we then present a simple statistical method to evaluate whether at least one of a set of apparently diagnostic characters is below the frequency cut-off. This method allows testing of the strength of the evidence for species distinctness and is readily applicable to empirical studies.     

Ant species differences determined by epistasis between brood and worker genomes

Epistasis arising from physiological interactions between gene products often contributes to species differences, particularly those involved in reproductive isolation. In social organisms, phenotypes are influenced by the genotypes of multiple interacting individuals. In theory, social interactions can give rise to an additional type of epistasis between the genomes of social partners that can contribute to species differences. Using a full-factorial cross-fostering design with three species of closely related Temnothorax ants, I found that adult worker size was determined by an interaction between the genotypes of developing brood and care-giving workers, i.e. intergenomic epistasis. Such intergenomic social epistasis provides a strong signature of coevolution between social partners. These results demonstrate that just as physiologically interacting genes coevolve, diverge, and contribute to species differences, so do socially interacting genes. Coevolution and conflict between social partners, especially relatives such as parents and offspring, has long been recognized as having widespread evolutionary effects. This coevolutionary process may often result in coevolved socially-interacting gene complexes that contribute to species differences.     

Identifying biochemical phenotypic differences between cryptic species

Molecular genetic methods can distinguish divergent evolutionary lineages in what previously appeared to be single species, but it is not always clear what functional differences exist between such cryptic species. We used a metabolomic approach to profile biochemical phenotype (metabotype) differences between two putative cryptic species of the earthworm Lumbricus rubellus. There were no straightforward metabolite biomarkers of lineage, i.e. no metabolites that were always at higher concentration in one lineage. Multivariate methods, however, identified a small number of metabolites that together helped distinguish the lineages, including uncommon metabolites such as Nε-trimethyllysine, which is not usually found at high concentrations. This approach could be useful for characterizing functional trait differences, especially as it is applicable to essentially any species group, irrespective of its genome sequencing status.     

Linking grazing response of species abundance to functional traits in the Tibetan alpine meadow

Whether grazing response of species abundance can be predicted by plant functional trait remains a challenge untested in a specific ecosystem. We measured species abundance and relevant functional traits for 30 common component species in a moderately grazed and a control community over 3 years (2005, 2006 and 2007) in a Kobresia capillifolia dominated alpine meadow on the Tibetan Plateau. Our objective was to examine species response to grazing and to test the relationship between the response of species abundance and functional traits in grazed and control habitats. Our results showed: i) in terms of species relative abundance and saturated height (the maximum height of an adult individual), most component species significantly decrease in response to moderate grazing and the effect differed among species and between functional groups. ii) The response of species abundance was significantly negatively correlated with saturated height, but not correlated with specific leaf area or seed size. We concluded that the response direction of species abundance to grazing can be predicted by functional traits, whereas it is a weak predictor of the extent of grazing response in species abundance.