Indirect effects of black spruce (Picea mariana) cover on community structure and function in sheep laurel (Kalmia angustifolia) dominated heath of eastern Canada

Recent studies on phenotypic plasticity of plant traits indicate that within-species variation in litter quality may be a significant factor that feeds back on litter decomposition and nutrient cycling rates at the stand level. These findings may be especially significant for understanding biodiversity-stability relationships in species-poor ecosystems that have little functional redundancy among primary producers. We tested the null hypothesis that black spruce and Kalmia were functional equivalents with respect to their structuring roles of subordinate vegetation and their influence on site biogeochemistry. The purpose of the study was to determine the degree to which forest cover exerts top-down control on community structure and function of Kalmia-black spruce communities. This community type dominates much of the forest understory and unforested heathlands in Atlantic Canada. We intensively studied a representative stand of Kalmia heath in Terra Nova National Park in eastern Newfoundland. Thirty-two 0.5 m × 0.5 m sample plots were randomly distributed among five transects bisecting gradients in dominance of black spruce and Kalmia. Light levels, species composition, vascular plant cover and soil respiration rate were determined for each plot. Tissue samples of litter, mature and current year leaves of Kalmia were collected and analyzed for nutrient status. Herbaceous species richness and cover peaked at intermediate light levels. Kalmia foliar N concentration and above-ground biomass increased with increasing shade. Soil respiration rates were strongly related to the light gradient and increased with increasing quality of Kalmia litter inputs. Our data indicate that Kalmias vigour and foliar nitrogen concentrations are greater under black spruce canopy as opposed to heath condition and that the shaded phenotype has relatively benign feedbacks on soil productivity compared to the open-habitat phenotype. In the absence of functional diversity at the species level in these species-poor habitats, phenotypic plasticity in Kalmia appears to be an important dimension of the biodiversity-stability relationship in these communities since our data suggest that this species has the potential either to inhibit or facilitate carbon cycling and the pathway is strongly linked to the presence or absence of overstory cover. The role of forest regeneration as an indirect control of forest soil processes such as carbon and nitrogen cycling in this ecosystem is discussed.     

The effects of aperiodic desiccation on the diversity of benthic desmid assemblages in a lowland peat bog

The lowland minerotrophic “Swamp” peat bog, located in the Czech Republic, is one of the most important European sites of desmidiacean diversity. The hydrological regime of the bog is driven by the water level of a nearby ancient manmade pond. Therefore, the bog experiences severe aperiodic drying events related to the pond draining. In this study, we investigated the drought-related response of the benthic desmid assemblages of the bog. The samples were taken bimonthly from 12th May 2008 to 19th May 2010, including the 8-month drying out period between October 2008 and June 2009. In addition to the species frequency data, morphometric methods were used to analyse the disparity, morphological turnover and surface-to-volume (S:V) ratios of Desmidiales. The drying event influenced the species composition, biovolume and S:V ratio data of the more species-rich pool, but its influence was less conspicuous in the species-poor acidic site. Accordingly, the species-rich site had generally higher species or morphological turnover between successive samples. The indicators based on the morphometric data were generally more sensitive than the species data. Therefore, we propose that the biovolume, S:V ratios or disparity measures of desmidiacean assemblages might be of benefit for future studies of peat bog microphytobenthos. Desmid assemblages at both sites recovered rapidly following the re-wetting of the bog, and they attained the pre-disturbance diversity, species composition and disparity values. We conclude that the drying event of the bog did not irreversibly influence its valuable desmid assemblages.     

The age of Calluna stands moderates post‐fire regeneration rate and trends in northern Calluna heathlands

Questions: Does stand age influence the direction and rate of post‐fire successional dynamics in coastal Calluna heaths and can old degraded heath vegetation be restored through reintroduction of fire? Location: Coastal heaths in the Tarva archipelago, central Norway. Methods: We investigated revegetation dynamics after experimental fires set in young (8 years since last fire) and old (>50 years since last fire) grazed heath stands. A repeated measures design was used, with floristic data recorded in permanent plots in the post‐fire successions (n=12) over a 7‐year period. The data were analysed using multivariate ordination techniques (PCA, RDA and PRC) and mixed effects models. Results: The age of Calluna stands strongly influenced post‐fire succession, different trends due to age explained 10.4% of variation in floristic data. Young heath showed faster succession towards pre‐fire community composition than old heath, and this could partially be explained by succession‐related factors: young heath had lower cover of mosses and lichens in the pre‐burned vegetation, and lower cover of litter early in succession. Young heath had a less pronounced overall community response to fire than old heath. Vegetative regeneration of C. vulgaris was absent in both old and young heath, but Calluna still re‐established as the dominant species within 5–7 years in both young and old stands. Regeneration dynamics were also affected by habitat conditions, different trends due to habitat explained 6% of variation. Conclusions: Our study demonstrates that old stands do develop characteristic heathland vegetation and structure after fire, and while potential invasives into the system such as trees and rhizomatous species are present, they do not impair Calluna regeneration or vegetation development towards the target heathland community composition and structure. Further, as our young stands are only in their second fire rotation after restoration, we suggest that characteristic dynamics of managed heathlands can re‐establish relatively rapidly, even in severely degenerated sites (>50 years since last fire). Site‐specific factors also need to be considered. We conclude that there is restoration potential in old heaths, despite slow dynamics in the first rotation.     

Markov models and initial floristic composition in heathland vegetation dynamics

A simple Markov model is used to test the hypothesis that the floristic composition of vegetation colonizing bare ground immediately after burning is the major factor influencing post-fire development in heathland vegetation. Data are taken from stands of different ages at time of burning in a species-rich Calluna-Arctostaphylos heath in NE Scotland. It is shown that variation in the initial floristic composition of the stands is not, in itself, sufficient to produce model simulations which match observed trends, although altering the probabilities of transition from bare ground to other states allows more successful simulations. The model supports the hypothesis that stand age before burning influences the post-fire development through the process of colonization of bare ground. After a very severe fire post-fire development may initially depend on the formation of a moss cover, although this requires further study. It is concluded that simple Markov models can provide the basis for examining successional processes when used in a comparative way.Nomenclature follows Tutin et al. (1964–80) for vascular plants and Smith (1978) for mosses.We thank Prof. C. H. Gimingham for comments on the draft, and J. M. M. Humphrey, P. Marren and the Nature Conservancy Council for permission to work on the Muir of Dinnet.     

Ecological Succession of a Relict Central European Peat Bog and Variability of Its Insect Biodiversity

The isolated habitat of the ervené Blato bog (South Bohemia, Czech Republic) and its relict insect fauna have been the subject of long-term monitoring. The species composition and abundance of Lepidoptera (light traps) and Coleoptera (pitfall traps) were monitored for 4 years (1994–1997) simultaneously on two sites – in the edaphic climax pine forest and in wetland successional habitats. The method of statistical evaluation by RDA and CCA ordination, representing the habitat preference of species of Coleoptera (Carabidae only) and Lepidoptera (all nocturnal phototactic taxa) between the edaphic climax forest and succession stages, was used. All categories of the peatland taxa (tyrphobiontic, tyrphophilous and tyrphoneutral species) were analysed. Ten highly stenotopic tyrphobiontic species and 23 tyrphophilous species of Lepidoptera (out of 487) were most characteristic of the bog habitat. Only two tyrphophilous carabid species (out of 20) were characteristic of the bog. The most important relict species (tyrphobionts) of Lepidoptera are most diverse and abundant within the successional habitats and in the open wet forest. The relict fauna of the closed climax pine forest is much less diverse and composed mostly of abundant tyrphophilous and tyrphoneutral forest species. Preservation or restoration of sufficiently constant hydrological conditions, which prevents formation of the closed forest, is the basic management for habitat conservation of all relict tyrphobiontic species of the ervené Blato bog and similar peat land habitat islands. The peat bog is a unified complex system of specific diverse and relict taxa. The most specific taxa are tyrphobiontic Lepidoptera, but a number of other vulnerable tyrphophilous and tyrphoneutral insects are associated with the peat bog as well.     

Biodiversity influences invasion success of a facultative epiphytic seaweed in a marine forest

The biotic resistance hypothesis predicts that more diverse communities should have greater resistance to invasions than species-poor communities. However for facultative and obligate epiphytic invaders a high native species richness, abundance and community complexity might provide more resources for the invader to thrive to. We conducted surveys across space and time to test for the influence of native algal species abundance and richness on the abundance of the invasive facultative epiphytic filamentous alga Lophocladia lallemandii in a Mediterranean Cystoseira balearica seaweed forest. By removing different functional groups of algae, we also tested whether these relationships were dependent on the complexity and abundance of the native algal community. When invasion was first detected, Lophocladia abundance was positively related to species richness, but the correlation became negative after two years of invasion. Similarly, a negative relationship was also observed across sites. The removal experiment revealed that more complex native communities were more heavily invaded, where also a positive relationship was found between native algal richness and Lophocladia, independently of the native algal abundance. Our observational and experimental data show that, at early stages of invasion, species-rich seaweed forests are not more resistant to invasion than species-poor communities. Higher richness of native algal species may increase resource availability (i.e. substrate) for invader establishment, thus facilitating invasion. After the initial invasion stage, native species richness decreases with time since invasion, suggesting negative impacts of invasive species on native biodiversity.     

The use of rotifer communities to provide a preliminary national classification of small water bodies in England

The rotifer species present in samples from 180 small water bodies, or ponds, in England were identified and listed for each pond. These communities were used to provide a system of classification of ponds applicable on a national basis, using the program TWINSPAN. Sites were also ordinated using DECORANA to identify major trends within the data set. The environmental factors most highly correlated with rotifer community were: (a) conductivity (P<0.001), (b) pH (P<0.001) and (c) permanence of water body (P<0.001). Out of 167 species, 32 were acting as indicators. It is suggested that a pond classification could be based on these species alone.     

Diversity and stability in garrigue ecosystems after fire

Summary Vegetation dynamics after fire was studied in six communities in Bas-Languedoc (Southern France). 47 plots were observed by means of a permanent transect for ten years.In the first part, we describe floristic richness, species fugacity and the way by which, the terminal community (as defined by the last observation) appears. The dynamics of all these communities follows a simple and general model: floristic richness reaches its maximum during the first two years after a fire, then decreases and becomes stable. Fugacity follows a similar model, whereas the mergence of the terminal community is rapid: one year after fire 70% of the plots have already acquired 75% of the species of the terminal community. There is no succession (in the general sense of the word), but a progressive reappearance of the species belonging to the original community.In the second part, we study floristic similarities between our plots and corresponding associations as described in literature. It appears that after a fire the floristic diversity of the landscape remains high; while the communities rapidly reach a relative maturity.In the study area fire seems to be a rather superficial phenomenon; it does not lead to an important modification of the community dynamics, because probably the most frequent species in Bas-Languedoc developed adaptations to withstand fire.     

Interactions between annuals and woody perennials in a Western Australian nature reserve

Abstract. We studied the interactions between woody perennial species and native and non-native annual species in a number of vegetation types within a nature reserve in the Western Australian wheatbelt. In particular, we examined the responses of annuals to perennial canopy removal, fire, soil disturbance and nutrient additions, and the effects of removal of annuals on perennial seedling regeneration. Experimental shrub removal significantly increased the abundance of annuals in a dense shrubland dominated by Allocasuarina campestris, but had no effect in a more open species-rich sandplain heath. Soil disturbance and nutrient addition in the heath area had no significant influence on annual abundance until three years after treatment. Fire had no clear effect on annual abundance in the heath within the reserve, but promoted a large increase in non-native species within an adjacent roadverge. A pattern of increased soil nutrient levels was accompanied by greatly increased non-native annual abundance beneath individual trees of Santalum spicatum. Exploratory laboratory bioassay experiments indicated that several woody perennials produced leachates that were capable of reducing the germination or growth of the introduced grass Avena fatua, indicating that allelopathy may be an important component of the interaction between the annual and perennial components. Within a woodland community, fire temporarily reduced the abundance of annual species and increased the establishment of perennial seedlings. Field experiments showed that annuals significantly reduced the survival of seedlings of the shrub Allocasuarina campestris. Our results indicate that intact native vegetation canopies effectively prevent invasion by non-native annuals, and that regeneration by native perennials is likely to be inhibited by the presence of an abundant annual cover.     

Communities containing Juncus squarrosus in upper Teesdale,England

Summary Plant communities containing Juncus squarrosus were analysed in a 100 km² area at the head of Teesdale in the north Pennines. The method of successive approximation was applied, using quadrats of 1/4m². Five noda were recognised, and their relation to noda previously described from the Scottish Highlands is discussed.The species-rich Juncus-Nardus nodum on irrigated gley soils, and the Juncus-Festuca rubra-Mnium punctatum nodum on flushed peat, contain many hydrophilic species and together constitute an association called species-rich Juncetum squarrosi sub-alpinum. The species-poor Juncus-Nardus nodum occurs on gley soils poorer in bases, and is probably a facies of species-poor Nardetum sub-alpinum. The remaining two noda, the Juncus-Vaccinium myrtillus and the Juncus-Carex nigra-Polytrichum commune, are found on acid, humus-rich soils, being differentiated by freedom of drainage. They contain little Nardus, but constitute an association called species-poor Juncetum squarrosi sub-alpinum. This occupies a considerable area in upper Teesdale, but differs from similar communities described in the Carneddau of north Wales and in the Scottish Highlands in that Agrostis spp. and Anthoxanthum odoratum are scarce. This is perhaps due to long-continued grazing by sheep.     

Length of burning rotation and community composition in high-level Calluna-Eriophorum bog in N England

Studies on a long-term experiment on Calluna-Eriophorum bog indicate that the length of time between management fires affects the post-fire development. A short rotation (every 10 years) results in increased dominance by Eriophorum spp. while a long rotation (every 20 years) results in greater abundance of Calluna vulgaris after fire. Variations in post-fires species abundances are related to pre-fire stand composition coupled with the patchy distribution of regeneration centers of rhizomatous species. The usefulness of burning this vegetation type in management for herbivore grazing is questioned.Nomenclature follows Tutin et al. (1964 80) for vascular plants. Smith (1978) for mosses, Watson (1981) for liverworts and Hawksworth, James & Coppins (1980) for lichens. Following common practice. Calluna vulgaris is referred to simply as Calluna. Acknowledgements. I thank M. Raves and the Nature Conservancy Council for permission to work on Moor House National Nature Reserve and for the use of data from Reserve Records, V. J. Hobbs for field assistance, and Prof. C. H. Gimingham for help and advice during the study. I also thank all who worked at Moor House for creating a pleasant working environment. The long-term experiment was planned by Dr R. J. Elliot and analyzed initially by Dr G. I. Forrest.     

Population structure and dynamics of a neotropical palm in fire-impacted fragments of the Brazilian Atlantic Forest

The demography of the acaulescent palm Attalea humilis was studied from 1996 to 1999 in three forest fragments (1.6, 6.4, and 9.9ha) in southeastern Brazil (22°30–22°33 S, 42°15–42°19 W). A human-induced fire damaged the populations in August–September 1997. Palm density was higher in the medium-sized fragment, where it was concentrated in a single disturbed sector. Population ontogenetic stage structure was dependent on fragment and year, but in general later stages were more abundant than initial ones. Population finite growth rates projected from linear, stage-structured demographic models indicated that the populations were increasing before fire. Population growth rate in the smallest fragment was larger due to a single recruitment event, being most sensitive to changes in survival and growth of seedlings, virgins and the immature, and relatively insensitive to changes in shrinkage and fecundity. Observed growth rates for the entire study period showed that populations remained in equilibrium after fire, but the population in the medium-sized fragment showed a declining tendency. Population structure was resilient to fire, which caused a transitory concentration of individuals in the medium-sized stage. The self-sustainability of the population in the largest, most preserved fragment suggests that habitat-protection measures may suffice to ensure the persistence of populations in small fragments. However, population fluctuations and exclusion from hyper-disturbed sectors in the smallest fragment suggest that reduced fragment area, habitat degradation and human-induced fires can interact to threaten the persistence of even abundant, preadapted to fire, species like A. humilis.     

Contrasting changes in vegetation structure and diversity with time since fire in two Australian Mediterranean‐climate plant communities

Mallee‐heath and mallee communities occur in a mosaic across large areas of south‐western Australia, in topographically subdued and fire‐prone landscapes. Consequently, it could be expected that these communities would have historically experienced similar fire regimes, and would respond similarly to variation in aspects of the fire regime. We studied the response of mallee‐heath and mallee to time since the last fire, measuring species density, species–area relationships, diversity indices and vegetation structure. Floristic responses to time since fire accorded with the initial floristic composition model of plant succession, with declining species density and Shannon diversity with age in mallee‐heath. Mallee‐heath exhibited structural senescence when > approximately 45–55 years since fire, with increasing standing dead vegetation, bare ground and stagnating or declining size in sprouting Eucalyptus spp. Mallee showed no such evidence of senescence, and indeed continued to increase in stature beyond the mean fire interval but without the compositional change required to provide support for the relay floristic model of plant succession. These results indicate that mallee‐heath is a fire maintained community and as such is reliant upon periodic burning to maintain diversity and vigour. Mallee, in contrast, is modified but not maintained by fire (at least over the period of time since fire examined) and hence is less susceptible to fire interval effects. Indeed, structural attributes likely to be significant for fauna habitat and carbon sequestration continue to develop in mallee unburnt for 55 years or more. Different responses to time since fire will create challenges for management, particularly in fragmented landscapes where fire potentially interacts with other threatening processes.     

The systematics of Porphyra: character evolution in closely related species

Isozymes, vegetative and reproductive morphology, seasonality, vertical and geographic distributions and chromosomes were compared for six pairs of putative sibling species of Porphyra (P. abbottae/P. torta, P. fallax subsp. fallax/P. fallax subsp. conwayae, P. amplissima/P. cuneiformis, P. fucicola/P. leucostica, P. miniata/P. variegata, P. umbilicalis/P. umbilicalis) and among five species in a complex (P. brumalis, P. kurogii, P. linearis, P. pseudolinearis, and P. purpurea.) Geographic distribution and zymograms for certain proteins showed the greatest change between species pairs: only one pair of species had identical distributions, and most species pairs were disjunct; every species had a different allozyme for GOT-1, whereas all species had apparently identical proteins for phycoerythrin. Seasonality and habitat exhibited moderate differentiation: Northeast Pacific sibling species were characterized by a high intertidal winter species pairing with a mid intertidal spring species, whereas all but one of the other species pairs exhibited nearly identical vertical distributions and seasonalities. There were few changes in morphology: most species pairs had essentially identical morphologies and coloration and the same arrangement of reproductive cells. Chromosome numbers and karyotypes were identical for species pairs and in the species complex. These results provide evidence for different rates of evolution of different characters in the genus Porphyra.     

Early post-fire vegetation regeneration in <Emphasis Type="Italic">Larix kaempferi</Emphasis> artificial forests with an undergrowth of <Emphasis Type="Italic">Sasa senanensis</Emphasis>

Post-fire vegetation regeneration was studied for a 6-year period in a 13-year-old-artificial forest consisting of Larix kaempferi with a dense undergrowth of Sasa senanensis. The study site was classified into three fire severity categories according to the degree of Sasa senanensis scorching, that is, a high-severity category, a mid-severity category, and a low-severity category. Study plots were established in areas which fitted the criteria for each category, and in nearby unburned sites. A total of 41 woody species were newly emerged during the 6-year study period in the burned and unburned plots. Only a few seedlings and resprouts emerged in the unburned plots, while many seedlings emerged in the high-severity plots from the first year after fire onward. A high-severity fire that burns the rhizomes of Sasa is necessary for the vegetation recovery by germination of seed. Whereas the establishment of seedlings was restricted to a few years after fire, the regeneration through resprouting continued into the last year of observation. The survival time of resprouts was longer than that of seedlings, and the survival time of shade-tolerant species was longer than that of shade-intolerant species. In contrast, shade-intolerant species grew more rapidly than shade-tolerant species. The plants ability to exceed the maximum height of the Sasa before the bamboo recovers can be critical to the survival of shade-intolerant species. Because resprouts have a stronger resistance to the shade of Sasa than seedlings, the resprouts of shade-tolerant species play a major role in the re-establishment of woody species after fire in sites with considerable Sasa ground-cover.     

Effets de la mise en eau du réservoir Desaulniers (Territoire de la Baie de James) sur le zooplancton d'une rivière et d'une tourbière reticulée

This paper presents the results of an investigation which lasted three years, on the effects of impounding a river on its zooplankton community. Forty zooplankton samples were collected from the Desaulniers River and from a string bog (53°35N, 77°35W) in 1976 before impoundment. Sampling was also conducted in 1977 (331 samples) and 1978 (153 samples) during and after impoundment. The population dynamics of zooplankton show different patterns in 1977 and 1978. The main impacts of impoundment on the river and the bog communities are the increase of the population density, particulary of Rotifera and Copepoda, the increase of the number of species of Rotifera and the increase of the diversity indices based on numbers and biomass. These changes are observed at each sampling station. The total biomass increases in lotic stations but decreases in the string bog because the density and the biomass of Cladocera increases in the river but drops in the bog. Results are compared to those obtained in natural lakes and bog of the same area and to those published in other studies.

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Effets de la mise en eau du réservoir Desaulniers (Territoire de la Baie de James) sur le zooplancton d'une rivière et d'une tourbière reticulée

     

Testing for non-randomness in sizes and habitats of West Indian lizards: choice of species pool affects conclusions from null models

Summary Several null models are proposed for testing whether size or habitat differences in West IndianAnolis lizards are greater than expected by chance. The models differ primarily in choice of the pool from which species are sampled to form random communities. Regardless of choice of pool, size differences in the Lesser Antilles are greater than null models predict; the pool using species on the known source (Puerto Rico) gives a greater variance in ratios but about the same mean ratio (for males), or a greater mean ratio (for females), compared with the pool composed of species on the islands being tested (the Stronget al., 1979, Galápagos procedure). On satellite islands of the Greater Antilles, sizes do not differ more than expected from null models. Pools composed of mainland-source species give null communities with more small or more large ratios than those composed of island species, depending upon whether four-species islands are included or excluded, respectively. Colwell and Winkler's unmodified Narcissus hypothesis is contradicted by these results in procedures where species not likely to be able to occur on small islands are included in the species pool. Using the most biologically reasonable, but not other, choices of source pool, species on satellite islands of the Greater Antilles differ more in structural habitat then expected by chance. In contrast to some of the results on size, here mainland-source pools are more likely to produce a statistically significant difference between real and random communities, as predicted by the Narcissus hypothesis. However, exclusion of structural habitat categories not found on satellite islands is necessary to achieve this significance.     

Predator-prey ratios in communities of freshwater invertebrates: the role of enemy free space

SUMMARY.

• 1 The ratio: number of predator species/number of prey species is reviewed using comprehensive faunal lists for a range of freshwater habitats in Britain and North America. Prey species are defined as detritivores, herbivores and fungivores; predators eat metazoan animals as the main component of their diet. Our data refer only to invertebrates.
• 2 The numbers of predators and prey species are apparently very closely correlated in freshwater communities (r=0.84, In transformed data), with an average ratio of predators to prey of 0.36. The average ratio of predators to prey changes from 0.48 in small (species-poor) collections to 0.29 in large (species-rich) collections.
• 3 We suggest that an approximately constant ratio of predators to prey may be generated by: (a) the number of predator species being a function of the number of broad classes or kinds of prey; and (b) the number of prey species being constrained by competition between prey for ‘enemy free space’, i.e. species that are too similar are unable to coexist with shared predators.