Was total primary production in the western Wadden Sea stimulated by nitrogen loading?

Borum and Sand-Jensen (1996) described empirical relationships between nitrogen (N) loadings from land and total (benthic + pelagic) primary production rates in shallow coastal marine waters. We applied these relationships to N loadings of the western Wadden Sea system, and compared the production estimates with actually observed primary production rates of autotrophic components (phytoplankton, microphytobenthos, macroalgae and seagrasses) for those years for which field data were available. During the 1980s and early 1990s, primary production values appear in good agreement with those derived from the empirical relationships. During the 1960s and early 1970s, however, these relationships substantially overestimated the total primary production in the western Wadden Sea. Based on ambient nutrient concentrations and the Redfield ratio, production in that period was considered not to be limited by N but by phosphorus (P) during most of the time. It is concluded that primary production is not invariably stimulated by N loading from land. If other factors (i.e. additional nutrient sources, N:P ratios, internal nutrient dynamics and co-limiting effects of nutrients and light) are not taken into account, management regulations that are targeted at diminishing the effects of eutrophication hold the risk of seriously under- or overestimating nutrient reductions that are thought necessary to achieve their goals. Received: 30 November 1998 / Received in revised form: 12 July 1999 / Accepted: 15 July 1999     

Effects of livestock grazing on aboveground insect communities in semi-arid grasslands of southeastern Arizona

Despite the importance of invertebrates in grassland ecosystems, few studies have examined how grassland invertebrates have been impacted by disturbances in the southwestern United States. These grasslands may be particularly sensitive to one common disturbance, livestock grazing, because they have not recently evolved in the presence of large herds of bison, an important mammalian herbivore. This study examined how livestock grazing influenced vegetation-associated insect communities in southeastern Arizona. Insect abundance, richness, diversity, community composition, and key environmental variables were compared between sites on active cattle ranches and sites on a 3160 ha sanctuary that has not been grazed by cattle for over 25 years. Vegetation-associated insect communities were found to be sensitive to livestock grazing. Overall abundance of these insects was lower on grazed grasslands, and certain insect orders appeared to be negatively affected by livestock grazing; beetles were less rich, flies were less diverse, and Hymenoptera were less rich and diverse on grazed sites. Conversely, Hemiptera were more diverse on grazed sites. Species composition of vegetation-associated insect communities also differed and was significantly correlated with percent vegetation cover and number of shrubs. Insect species responsible for these differences were taxonomically diverse, and included herbivores and predators/parasites. When compared to other studies conducted in areas of the United States that fall within the historic range of bison, this study suggests that invertebrates in areas outside this range may be more sensitive to grazing pressure.     

Host plant species effects on arbuscular mycorrhizal fungal communities in tallgrass prairie

Symbiotic associations between plants and arbuscular mycorrhizal (AM) fungi are ubiquitous in many herbaceous plant communities and can have large effects on these communities and ecosystem processes. The extent of species-specificity between these plant and fungal symbionts in nature is poorly known, yet reciprocal effects of the composition of plant and soil microbe communities is an important assumption of recent theoretical models of plant community structure. In grassland ecosystems, host plant species may have an important role in determining development and sporulation of AM fungi and patterns of fungal species composition and diversity. In this study, the effects of five different host plant species [Poa pratensis L., Sporobolus heterolepis (A. Gray) A. Gray, Panicum virgatum L., Baptisia bracteata Muhl. ex Ell., Solidago missouriensis Nutt.] on spore communities of AM fungi in tallgrass prairie were examined. Spore abundances and species composition of fungal communities of soil samples collected from patches within tallgrass prairie were significantly influenced by the host plant species that dominated the patch. The AM fungal spore community associated with B. bracteata showed the highest species diversity and the fungi associated with Pa. virgatum showed the lowest diversity. Results from sorghum trap cultures using soil collected from under different host plant species showed differential sporulations of AM fungal species. In addition, a greenhouse study was conducted in which different host plant species were grown in similar tallgrass prairie soil. After 4 months of growth, AM fungal species composition was significantly different beneath each host species. These results strongly suggest that AM fungi show some degree of host-specificity and are not randomly distributed in tallgrass prairie. The demonstration that host plant species composition influences AM fungal species composition provides support for current feedback models predicting strong regulatory effects of soil communities on plant community structure. Differential responses of AM fungi to host plant species may also play an important role in the regulation of species composition and diversity in AM fungal communities. Received: 29 January 1999 / Accepted: 20 October 1999     

The relationship between productivity and multiple aspects of biodiversity in six grassland communities

Biodiversity is a multifaceted concept but most studies examining the association between the biodiversity of a community and its productivity focus only on species richness. Consequently, studies are needed to examine how other facets of biodiversity vary with productivity if we want to have a better understanding of the distribution of biodiversity across our planet. We evaluated how a number of biodiversity measures (species richness, evenness, dominance, rarity, Simpson’s diversity, and Shannon–Weiner diversity) varied across natural productivity gradients at 6 grassland sites in the continental US. Variation in productivity did not account for a substantial amount of variation in any measure of biodiversity at small spatial scales (≈1 m²) at most sites. When productivity accounted for substantial variation in biodiversity, different measures of biodiversity responded to productivity in different ways. For example, dominance changed in a U-shaped fashion along a productivity gradient whereas richness increased in an asymptotic fashion. Consequently, diversity indices, which account for both species richness and evenness, varied in a hump-shaped fashion along the productivity gradient. Our results highlight that an exclusive focus on the association between species richness and productivity provides an incomplete picture of how a community’s biodiversity is related to its functioning.     

Observational evidence that diversity may increase productivity in Mediterranean shrublands

This paper reports field observational evidence on the diversity-biomass production relationship from eastern Mediterranean shrublands dominated by Cistus salviifolius. The main results are: (1) plant diversity and ecosystem primary production are positively correlated in the Cistus shrublands, which are multigrowth form systems; (2) the taxonomic and growth form diversity in Cistus shrublands suggests that there is strong differentiation along the structural niche axis, and thus functional redundancy is weak; (3) the performance of the dominant Cistus in terms of biomass production did not affect the overall diversity-biomass production of the studied communities. Received: 18 October 1999 / Accepted: 13 April 2000     

Seagrass bed patchiness: effects on epifaunal communities in San Diego Bay, USA

Seagrass habitat structure influences epifaunal density, diversity, community composition and survival, but covariation of structural elements at multiple scales (e.g., shoot density or biomass per unit area, patch size, and patch configuration) can confound studies attempting to correlate habitat structure with ecological patterns and processes. In this study, we standardized simulated seagrass shoot density and bed area among artificial seagrass beds in San Diego Bay, California, USA to evaluate the singular effect of seagrass bed configuration (“patchiness”) on the density and diversity of seagrass epifauna. Artificial seagrass beds all were 1 m², but were composed of a single large patch (“continuous” treatment), four smaller patches (“patchy” treatment), or 16 very small patches (“very patchy” treatment). We allowed epifauna to colonize beds for 1 month, and then sampled beds monthly over the next 3 months. Effects of seagrass bed patchiness on total epifaunal density and species-specific densities were highly variable among sampling dates, and there was no general trend for the effects of fragmentation on epifaunal densities to be positive or negative. Epifaunal diversity (measured as Simpson's index of diversity) was highest in very patchy or patchy beds on two out of the three sampling dates. Very patchy beds exhibited the highest dissimilarity in community composition in the first two sampling periods (August and September), but patchy beds exhibited the highest dissimilarity in the third sampling period (October). Our results indicate that seagrass patch configuration affects patterns of epifaunal density, diversity, and community composition in the absence of covarying bed area or structural complexity, and that patchy seagrass beds may be no less valuable as a habitat than are continuous seagrass beds. The spatial pattern employed when harvesting or planting seagrass may influence epifaunal habitat use and should be a key consideration in restoration plans.     

The embryonic expression pattern of labial, posterior homeotic complex genes and the teashirt homologue in an apterygote insect

 During embryogenesis of the fruit fly, Drosophila melanogaster, the homeotic genes are required to specify proper cell fates along the anterior-posterior axis of the embryo. We cloned partial cDNAs of homologues of the Drosophila homeotic gene teashirt and five of the homeotic-complex (HOM-C) genes from the thysanuran insect, Thermobia domestica, and assayed their embryonic expression patterns. The HOM-C genes we examined were labial, Antennapedia, Ultrabithorax, abdominal-A and Abdominal-B. As the expression pattern of these HOM-C genes is largely conserved among insects and as Thermobia is a member of a phylogenetically basal order of insects, we were able to infer their ancestral expression patterns in insects. We compare the expression patterns of the Thermobia HOM-C genes with their expression in Drosophila and other insects and discuss the potential roles these genes may have played in insect evolution. Interestingly, the teashirt homologue shows greater variability between Thermobia and Drosophila than any of the HOM-C genes. In particular, teashirt is not expressed strongly in the Thermobia abdomen, unlike Drosophila teashirt. We propose that teashirt expression has expanded posteriorly in Drosophila and contributed to a homogenization of the Drosophila larval thorax and abdomen. Received: 23 July 1998 / Accepted: 1 November 1998     

Stimulation of grassland nitrogen cycling under carbon dioxide enrichment

 Nitrogen (N) limits plant growth in many terrestrial ecosystems, potentially constraining terrestrial ecosystem response to elevated CO₂. In this study, elevated CO₂ stimulated gross N mineralization and plant N uptake in two annual grasslands. In contrast to other studies that have invoked increased C input to soil as the mechanism altering soil N cycling in response to elevated CO₂, increased soil moisture, due to decreased plant transpiration in elevated CO₂, best explains the changes we observed. This study suggests that atmospheric CO₂ concentration may influence ecosystem biogeochemistry through plant control of soil moisture. Received: 18 December 1995 / Accepted: 19 June 1996     

Nitrogen and energy requirements of the short-tailed fruit bat (Carollia perspicillata): fruit bats are not nitrogen constrained

 Nitrogen (N) and energy (E) requirements were measured in adult Carollia perspicillata which were fed on four experimental diets. Bats ate 1.3–1.8 times their body mass ⋅ day^-1 and ingested 1339.5–1941.4 kJ ⋅ kg^-0.75 ⋅ day^-1. Despite a rapid transit time, dry matter digestibility and metabolizable E coefficient were high (83.3% and 82.4%, respectively), but true N digestibility was low (67.0%). Mass change was not correlated with E intake, indicating that bats adjusted their metabolic rate to maintain constant mass. Bats were able to maintain constant mass with digestible E intake as low as 1168.7 kJ ⋅ kg^-0.75 ⋅ day^-1 or 58.6 kJ ⋅ . Metabolic fecal N and endogenous urinary N losses were 0.87 mg N ⋅ g^-1 dry matter intake and 172.5 mg N ⋅ kg^-0.75 ⋅ day^-1, respectively, and bats required 442 mg N ⋅ kg^-0.75 ⋅ day^-1 (total nitrogen) or 292.8 mg N ⋅ kg^-0.75 ⋅ day^-1 (truly digestible nitrogen) for N balance. Based on E and N requirements and digestibilities, it was calculated that non-reproductive fruit bats were able to meet their N requirements without resorting to folivory and without over-ingesting energy. It was demonstrated that low metabolic fecal requirements allowed bats to survive on low-N diets. Accepted: 23 June 1996     

Characterization of terrestrial insect communities using quantified, Malaise-trapped Coleoptera

Abstract. 1. Weekly samples of Malaise trapped Coleoptera from regenerating shrubland and an adjacent mature, podocarp-broadleaved forest, in the North Island, New Zealand, were compared over one season. Three traps were monitored in each site.
2. Diversity indices did not group samples by site whereas the divisive polythetic classification analysis TWINSPAN showed that the two sites possessed clearly different communities. In the classification, site was of primary importance, with time of year, and trap position within site of secondary and tertiary importance respectively.
3. Samples were compared at different taxonomic levels and using different subsets of the database. Determination of the main families, morphotyped to species or species complex, was found to be sufficient to classify most samples to the appropriate community group.
4. The divisive classification procedure applied to four consecutive weekly Coleoptera samples over early summer, is suggested as a means of describing and identifying terrestrial arthropod communities characteristic of site and year of collection. This approach provides a potentially sensitive tool for monitoring terrestrial ecosystems.     

Effect of organic and inorganic nitrogen sources on endogenous polyamines and growth of ectomycorrhizal fungi in pure culture

 The abilities of three ectomycorrhizal fungi, Paxillus involutus, Suillus variegatus and Lactarius rufus, to utilize organic and inorganic nitrogen sources were determined by measuring the growth and endogenous free polyamines (putrescine, spermidine and spermine) of pure culture mycelium. Differences were found in the utilization of the nitrogen sources and in the polyamine concentrations between the fungal species and between isolates of L. rufus. All the fungi grew well on ammonium and on several amino acids. Endogenous polyamine levels varied with the nitrogen source. Spermidine was commonly the most abundant polyamine; however, more putrescine than spermidine was found in P. involutus growing on inorganic nitrogen or arginine. Low amounts of spermine were found in S. variegatus and some samples of L. rufus. None or only a trace of spermine was found in P. involutus mycelium. In all fungi, putrescine concentrations were higher with ammonium than with the nitrate treatment. The total nitrogen content of peat did not determine the ability of L. rufus strains isolated from peatland forest sites to utilize organic nitrogen. Accepted: 27 November 1998     

Comparison of volatile leaf compounds and herbivorous insect communities on three willow species

 Plants produce volatile compounds known to influence insect preferences for oviposition and feeding. To examine whether volatile leaf compounds are correlated with the herbivorous insect community, we analyzed volatile compounds in leaves from three co-occurring willow species, Salix serissaefolia, S. eriocarpa, and S. integra, and investigated their associated insect communities in 3 months across different years. The gas chromatographic profiles of volatile compounds were highly specific to each willow species and remained constant in the study months. In a comparison between the chemical composition of the volatile compounds and the taxon composition of the insect communities, dissimilarity patterns in chemical composition among the three willow species were very close to those in herbivorous insect communities. These findings indicate that willow leaves produce specific volatiles that are highly correlated with the community structure of herbivorous insects associated with them. Received: October 10, 2002 / Accepted: March 17, 2003     

Context-dependent effects of bluegill in experimental mesocosm communities

Most knowledge of direct and indirect effects of zooplanktivorous fish has come from studies in which a treatment with a zooplanktivore is compared to a fishless control. However, effects of a zooplanktivore may be different in the presence of other fish species because the other fish have direct and indirect effects that may alter the effects of the zooplanktivore in question. To test this hypothesis, we conducted a tank mesocosm experiment of 2×2 factorial design in which the presence and absence of bluegill (Lepomis macrochirus) were cross-classified with the presence and absence of a fish assemblage composed of western mosquitofish (Gambusia affinis) and common carp (Cyprinus carpio). The presence of bluegill decreased Daphnia, Ceriodaphnia, cyclopoid copepodids, calanoid copepodids, copepod nauplii, amphipods, gastropods, and notonectids. Daphnia, Ceriodaphnia, cyclopoid copepodids, copepod nauplii, gastropods, notonectids, Najas, and Chara were decreased and herbivorous rotifers, turbidity, chlorophyll a, total nitrogen and total phosphorus were increased in the presence of the fish assemblage. Significant bluegill×fish assemblage interaction effects were detected for Daphnia, Ceriodaphnia, cyclopoid copepodids, copepod nauplii, gastropods, and notonectids. Analysis of simple effects for these response variables revealed that all significant bluegill effects in the absence of the fish assemblage were not significant in the presence of the fish assemblage. Our results indicate that the effects of bluegill may be context dependent, or dependent upon the presence of other trophically similar fish species. Received: 3 November 1998 / Accepted: 24 September 1999     

Effects of below-ground insects, mycorrhizal fungi and soil fertility on the establishment of Vicia in grassland communities

 The effects of, and interactions between, insect root feeders, vesicular-arbuscular mycorrhizal fungi and soil fertility on the establishment, growth and reproduction of Vicia sativa and V. hirsuta (Fabaceae) were investigated in an early-successional grassland community. Seeds of both species were sown into plots where soil insecticide (Dursban 5G), soil fungicide (Rovral) and soil fertiliser (NPK) were applied in a factorial randomised block design. Fertiliser addition reduced growth, longevity and reproduction of both Vicia species, due to the commonly recorded increase in the competitive advantage of the non-nitrogen-fixing species when nitrogen is added to the plant community. However, in plots where fertiliser was not applied, a reduction in root feeders and mycorrhizal infection led to an increase in seedling establishment and fruit production of V. sativa, and to an increase in flower production for both Vicia species. The interaction between all three soil treatments explained much of the variation in growth and longevity of V. sativa. Plants grew larger and survived longer in plots where natural levels of mycorrhizal infection and root feeders were low compared with plots where all the treatments were applied. This suggests that, although soil nutrient availability was a strong determinant of the performance of these two leguminous species, at natural levels of soil fertility biotic factors acting in the soil, such as mycorrhizal fungi and soil-dwelling insects, were important in shaping the competitive interactions between the two Vicia species and the plant community. Our results indicate that non-additive interactions between ecological factors in the soil environment may strongly affect plant performance. Received: 18 July 1995 / Accepted: 14 August 1996     

Effects of non-native plants on the native insect community of Delaware

Due to the lack of a co-evolutionary history, the novel defenses presented by introduced plants may be insurmountable to many native insects. Accordingly, non-native plants are expected to support less insect biomass than native plants. Further, native insect specialists may be more affected by introduced plants than native generalist herbivores, resulting in decreased insect diversity on non-native plants due to the loss of specialists. To test these hypotheses, we used a common garden experiment to compare native insect biomass, species richness, and the proportion of native specialist to native generalist insects supported by 45 species of woody plants. Plants were classified into three groupings, with 10 replicates of each species: 15 species native to Delaware (Natives), 15 non-native species that were congeneric with a member of the Native group (Non-native Congeners), and 15 non-native species that did not have a congener present in the United States (Aliens). Native herbivorous insects were sampled in May, June, and July of 2004 and 2005. Overall, insect biomass was greater on Natives than Non-native Congeners and Aliens, but insect biomass varied unpredictably between congeneric pair members. Counter to expectations, Aliens held more insect biomass than did Non-native Congeners. There was no difference in species richness or the number of specialist and generalist species collected among the three plant groupings in either year, although our protocol was biased against sampling specialists. If these results generalize to other studies, loss of native insect biomass due to introduced plants may negatively affect higher trophic levels of the ecosystem.     

Interactions between goldenrod (Solidago altissima L.) and its insect herbivore (Trirhabda virgata) over the course of succession

Consumers can mediate the composition of plant communities and alter ecosystem processes. Although herbivores usually increase N availability in the short term, they might decrease it in the long term. I investigated the long-term effect of insect herbivores on leaf tissue quality and soil N availability in goldenrod (Solidago altissima) fields using two approaches: (1) I compared plots from which herbivores had been excluded for 17 years with adjacent plots that had experienced normal levels of herbivory, and (2) I examined a chronosequence of nine goldenrod fields representing three successional stages: early, middle, and late. These parallel approaches showed that, in the long term, herbivores decrease the quality of leaf litter and soil N availability in goldenrod fields. These long-term effects appear to compensate for various short-term effects that increase N availability in the soil (e.g., added frass, increased light penetration). Furthermore, herbivores decrease leaf litter quality and N availability by reducing the quality of leaf tissue within the same species. This pattern may result from insect herbivores preferentially grazing on plants with a high N content thereby increasing the amount of recalcitrant litter over the course of succession. Received: 4 May 1999 / Accepted: 24 September 1999     

The effects of reduced phosphorus and nitrogen loading on phytoplankton in Mondsee,Austria

Following restoration measures (ring canalization, treatment plant with phosphorus precipitation), phosphorus loading declined step-wise from 26.21 t year^–1 to 9.18 t year^–1 during the period 1979–1984 while P-retention increased from 48% to 78%. Phosphorus loading was poorely correlated with precipitation. Inorganic nitrogen load, largely NO₃– N, did not decline but was significantly correlated with precipitation (r = 0.95) throughout the investigation period (1978–1989).Total phosphorus loading reached acceptable levels in 1984 when compared to critical loading calculated according to Vollenweider (1976). Phosphorus input to the lake has remained at these levels in recent years.Average annual chlorophyll-a concentrations and biovolume of phytoplankton in the top 20 m layer of the lake decreased, in correspondence with the respective phosphorus concentrations, from eutrophic to mesotrophic levels. The decline was accompanied by a drastic reduction of blue-green algal populations, and especially of Oscillatoria rubescens D.C..     

Role of brown bears (Ursus arctos) in the flow of marine nitrogen into a terrestrial ecosystem

We quantified the amount, spatial distribution, and importance of salmon (Oncorhynchus spp.)-derived nitrogen (N) by brown bears (Ursus arctos) on the Kenai Peninsula, Alaska. We tested and confirmed the hypothesis that the stable isotope signature (δ¹⁵N) of N in foliage of white spruce (Picea glauca) was inversely proportional to the distance from salmon-spawning streams (r=–0.99 and P<0.05 in two separate watersheds). Locations of radio-collared brown bears, relative to their distance from a stream, were highly correlated with δ¹⁵N depletion of foliage across the same gradient (r=–0.98 and –0.96 and P<0.05 in the same two separate watersheds). Mean rates of redistribution of salmon-derived N by adult female brown bears were 37.2±2.9 kg/year per bear (range 23.1–56.3), of which 96% (35.7±2.7 kg/year per bear) was excreted in urine, 3% (1.1±0.1 kg/year per bear) was excreted in feces, and <1% (0.3± 0.1 kg/year per bear) was retained in the body. On an area basis, salmon-N redistribution rates were as high as 5.1±0.7 mg/m² per year per bear within 500 m of the stream but dropped off greatly with increasing distance. We estimated that 15.5–17.8% of the total N in spruce foliage within 500 m of the stream was derived from salmon. Of that, bears had distributed 83–84%. Thus, brown bears can be an important vector of salmon-derived N into riparian ecosystems, but their effects are highly variable spatially and a function of bear density. Received: 11 February 1999 / Accepted: 7 July 1999