Colonization and early succession on anthropogenic soils

Abstract. This paper reports on vegetation development on permanent experimental plots during five years of succession. Nine (1 m²) plots were filled with three typical substrates from man-made habitats of urban and industrial areas in the region of Berlin. The three substrates (a commercial ‘topsoil’, a ruderal ‘landfill’ soil and a sandy soil), differ in organic matter and nutrient contents. Relevés of species composition and percent cover of each species present were made monthly during the growing season from the start of vegetation development. This paper describes the different successional pathways on topsoil and ruderal soil and the colonization process on sandy soil. On topsoil, ruderal annuals are dominant in the first year and are replaced by short-lived perennials from the second year. Those species were replaced by long-lived perennial herbs (Ballota nigra or Urtica dioica) from the third year of succession onwards. On the ruderal land-fill soil the early successional stages are less sharp and the perennial Solidago canadensis is able to dominate within one year after the succession was initiated. On sandy soil there is still an ongoing colonization process, where pioneer tree species like Betula pendula and Populus nigra play a main role. The importance of ‘initial floristic composition’, the role of substrate for community structure and the peculiarities of successional sequences on anthropogenic soils in the context of primary and secondary successions are discussed.     

The effect of prescribed burning on plant rarity in a temperate forest

Rare species can play important functional roles, but human‐induced changes to disturbance regimes, such as fire, can inadvertently affect these species. We examined the influence of prescribed burns on the recruitment and diversity of plant species within a temperate forest in southeastern Australia, with a focus on species that were rare prior to burning. Floristic composition was compared among plots in landscapes before and after treatment with prescribed burns differing in the extent of area burnt and season of burn (before–after, control‐impact design). Floristic surveys were conducted before burns, at the end of a decade of drought, and 3 years postburn. We quantified the effect of prescribed burns on species grouped by their frequency within the landscape before burning (common, less common, and rare) and their life‐form attributes (woody perennials, perennial herbs or geophytes, and annual herbs). Burn treatment influenced the response of rare species. In spring‐burn plots, the recruitment of rare annual herbs was promoted, differentiating this treatment from both autumn‐burn and unburnt plots. In autumn‐burn plots, richness of rare species increased across all life‐form groups, although composition remained statistically similar to control plots. Richness of rare woody perennials increased in control plots. For all other life‐form and frequency groups, the floristic composition of landscapes changed between survey years, but there was no effect of burn treatment, suggesting a likely effect of rainfall on species recruitment. A prescribed burn can increase the occurrence of rare species in a landscape, but burn characteristics can affect the promotion of different life‐form groups and thus affect functional diversity. Drought‐breaking rain likely had an overarching effect on floristic composition during our study, highlighting that weather can play a greater role in influencing recruitment and diversity in plant communities than a prescribed burn.     

Spontaneous vegetation succession in human‐disturbed habitats: A pattern across seres

Abstract. Vegetation samples from 15 successional seres in various disturbed habitats in the western part of the Czech Republic were analysed to detect possible trends. For particular seres, data on species cover were available from the onset to 10–76 yr of succession. All seres started on bare ground. Species which attained at least 1% cover in any sere in any year were used as input data for Canonical Correspondence Analysis, assessing the effect of time as the environmental variable, for Detrended Correspondence Analysis and TWINSPAN classification. Two distinct groups ofseres were distinguished: ‘ruderal’, occurring in agricultural, industrial or urban landscapes altered by men, usually on fertile sites; and ‘non‐ruderul’, occurring in less altered, mostly forested landscapes, usually on acid, nutrient‐poor and wetter soils. The former type of succession starts with ruderal annuals, being followed by ruderal perennials. In the latter case non‐ruderal clonal perennials prevail from the onset of succession. The landscape frame is emphasized, beside site environmental conditions, as influencing the type of succession. The character of species attaining dominance in succession, participation of dominant woody plants and the character of late successional stages, i.e. features important from the point of view of potential restoration of human‐disturbed habitats, are discussed.     

Spatio‐temporal variation in the effect of herbaceous layer on woody seedling survival in a Chilean mediterranean ecosystem

Questions: What is the effect of herbaceous layer on seedling establishment of three woody pioneer species in open areas of central Chile under a semi‐arid mediterranean climate? How do inter‐annual and habitat conditions (slope aspect) modulate this effect? Under high stress conditions such as the drier year and habitat (north‐facing slope) do herbs reach low abundance and have neutral effects on woody seedlings? Under medium stress conditions for these woody species, such as the wetter year and south‐facing slope, does the herbaceous layer reach greater abundance and have positive effects on woody seedlings due to increasing soil water content? Location: A watershed on the outskirts of Santiago, Chile, subjected to clearing of woody vegetation through firewood extraction and human‐set fires. Methods: In spring 2007, we set up 20 plots (3 m × 2 m). Half of each plot had herbs removed manually and by application of herbicide. In both halves of each plot, one seedling (8 months old) of each of the three native woody species (Colliguaya odorifera, Schinus polygamus and Quillaja saponaria) was planted and survival monitored subsequently. The experiment was repeated in two consecutive growing seasons (2007–2008 and 2008–2009) that differed significantly in total precipitation (152 and 256.5 mm, respectively), and replicated in two sites that differed in aspect and abiotic conditions: a moister south‐ and a drier north‐facing slope. Results: In the first and drier year, the herbaceous layer had low cover and no significant effect on seedling survival of woody species. During the second year, herbs had greater cover and a significant positive effect on spring survival of C. odorifera in the north‐facing slope, which was lost after summer. During this wetter year on the south‐facing slope, herb cover had a positive effect on survival of S. polygamus (mainly during summer). Conclusions: The role of mostly ruderal herbs on woody seedling establishment depended on the species, rainfall of the current year and slope aspect, and may be explained by soil moisture patterns. This suggests that the effect of ruderal herbs on woody seedlings shifts from neutral under high stress conditions produced by drought to positive under moderate stress conditions. Our results contribute to understand interactions between ruderal herbs and woody species under contrasting abiotic conditions. Therefore, control of the herbaceous layer may not be needed in restoration programmes for this region. Moreover, herbs may benefit restoration of woody cover in mesic habitats.     

Plant distribution and the temperature coefficient of metabolism

The spatial distribution of a plant species is limited by the range of climatic conditions to which the species can adapt. Temperature is one of the most significant determinants of plant distribution, but except for the effects of lethal limits, little is known about physiological changes in responses to differences in environmental temperature. In this study, temperature coefficients of non-photosynthetic metabolism have been determined in the normal environmental temperature range for selected annual and perennial plants. Distinct differences were found in the temperature coefficient of metabolism of woody perennial plants from high latitudes and high elevations and closely related low-latitude and low-elevation plants. Low-latitude and low-elevation woody perennials have Arrhenius temperature coefficients for metabolism that are larger than those for congeneric high-latitude and high-elevation plants. The Arrhenius temperature coefficient is not rapidly adapted to new environments. A simple function was developed relating Arrhenius temperature coefficient to latitude and elevation for accessions of three, woody, perennial species complexes of plants collected from a wide geographic range but grown in common gardens. Within these taxa, plants that experience broader ranges of temperature during growth in their native habitat have smaller temperature coefficients. Temperature coefficients also varied with growth stage or season. No similar relationship was found for annuals and herbaceous perennials. For the plants tested, Arrhenius temperature coefficients are high during early spring growth, but shift to lower values later in the season. The shift in Arrhenius temperature coefficients occurs early in the season for southern and low-elevation plants and progressively later for plants from further north or higher elevation. The changes in Arrhenius temperature coefficients result largely from increases in plant metabolic rates at lower temperatures while little change occurs in the rates at higher temperatures. Altering the temperature dependence of the control of metabolic rate is apparently an important means of response to climate change.     

Accumulation of buried seeds and establishment of ruderal therophytic communities in disturbed habitat,central Japan

The establishment and maintenance mechanisms of pioneer communities were investigated in ruderal habitats under two disturbance regimes, frequent and infrequent cutting sites. In the infrequent cutting sites, large perennials dominated through the year and inhibited the invasion of annuals, and the perennial community succeeded to forest stage if the cutting was stopped. In the frequent cutting sites, therophytic communities of winter and summer annuals alternated by season. Fresh seeds of both winter and summer annuals are dormant, but they have different germination times and thus can share the same sites in different seasons. Wind-dispersed biennials and large perennials have nondormant seeds and easily invade the sites; however, they are unable to mature to reproductive phase due to recurrent cuttings. The therophytic species, which can complete their life-cycle in a period between cuttings, accumulate seeds in the soils and are maintained by these buried seeds (seed bank annuals) during recurrent disturbances. The seed bank is compensation for the dispersal inefficiency of seed bank annuals. After abandonment of the frequent cutting sites, the buried seeds of seed bank annuals germinate and become the first-year pioneers. Thus, seed bank annuals are not invasive colonizers but are the remnants of the ruderal weed communities before abandonment.     

Perennial polypores as indicators of annual and red-listed polypores

Many polypores are specialized in their requirements for substrate and environment, and they have been suggested to indicate the continuity of coarse woody debris or naturalness of a forest stand. However, the use of polypores as indicators of conservation value is restricted by the temporally limited appearance of annual fruit bodies. We studied whether the species richness of perennial polypores (perennials) can be used to predict the species richness of annual or annual red-listed polypores (annuals). Our data included 1471 separate datasets (sample plots or larger inventoried areas) in different parts of Finland and Russian Karelia, ranging from the southern to northern boreal zone. At the large scale (the whole area) the number of perennials explained about 70% of the variation in the number of annuals, and about 67% in the number of red-listed annuals. A minimum set of 40–60 perennial occurrences gave a reliable estimate on the species richness of annuals, and 60–80 occurrences on the species richness of red-listed annuals. The richness of perennials predicted the richness of annuals and, in particular, richness of red-listed annuals, better than the size of inventoried area. According to our results, perennial polypores can be used as a surrogate for overall polypore species richness in natural and seminatural boreal forests, but the predictive power is weaker in managed forests. In addition, the relationship between the perennial and annual species seems to differ in different vegetation zones, management types and forest types. Due to this variation direct application of the indicator values derived from different vegetation zones and management or forest types are not recommended. Since perennials are easier to identify than annuals, detectable throughout the year, and have much smaller year-to-year variation, their use as an indicator group seems to offer advantages regarding the timing and cost-efficiency of inventories.     

Soil heterogeneity effects on canopy structure and composition during early succession

Early succession from annuals to herbaceous and woody perennials on abandoned fields and grasslands is marked by vegetation heterogeneity; i.e., variation in canopy composition and structure over the field through time. Soil resource heterogeneity could promote vegetation heterogeneity. We created soil resource patchiness on two newly ploughed fields by establishing two types (fertilized, unfertilized) of checkerboard plots with two, four, or eight subplots (trenched, untrenched) to test effects of soil fertility heterogeneity scale on vegetation heterogeneity during early succession. Canopy composition and structure were censused in years 2, 4 and 6. On both fields, soil resource heterogeneity did not affect canopy composition or structure. Differential abundance of dominant species, Ambrosia trifida on one field and Solidago altissima on the other, between fertilized and unfertilized plots led to decreased vegetation heterogeneity among subplots within plots and increased vegetation heterogeneity among plots. Soil enrichment promoted taller and layered, but species-poor, canopy. In general, soil fertility effects on abundance, expansion, and dominance of individuals of dominant species promoted variation in canopy composition and structure, or vegetation heterogeneity, among plots on the two fields.     

Impact of two wildfires on endemic granite outcrop vegetation in Western Australia

Abstract. In seasonally dry regions of the world fire is a recurring disturbance but little is known of how fire interacts with granite outcrop vegetation. We hypothesize that the floristic composition in granite vegetation, usually attributed to the edaphic environment, may also reflect the impact of disturbances such as fire. Dramatic differences in floristic composition and cover over 13 years and two fires were observed in vegetation on a Western Australian granite outcrop. This was very marked in the first year following the two fires, with annuals and geophytes showing the greatest turnover of species. Even among the perennial shrubs there was considerable turnover in a number of obligate seeders. After the first fire the number of species declined for woody perennials, herbaceous perennials and annuals, remained unchanged for perennial grasses and sedges, and varied with highest richness 4 yr after fire for geophytes. Demographic studies of two endemic woody obligate seeders and three endemic mallee eucalypt resprouters similarly showed dramatic differences within and between species in seedling recruitment following the two fires. Fire does have a significant impact on the floristic composition of semi‐arid granite outcrop vegetation communities. Studies on other granite outcrop systems are needed to test the generality of this conclusion.     

GERMINATION ECOPHYSIOLOGY OF HERBACEOUS PLANT SPECIES IN A TEMPERATE REGION

Germination phenology data have been collected from 75 winter annuals, 49 summer annuals, 28 monocarpic perennials, and 122 polycarpic perennials, and experimental investigations of dormancy breaking and germination requirements have been conducted on 56 winter annuals, 32 summer annuals, 18 monocarpic perennials, and 73 polycarpic perennials. The purpose of these studies was to determine if there are correlations between the dormancy breaking and germination requirements of seeds and the germination phenology, life cycle type, habitat requirements, range of geographical distribution, and phylogenetic relationships of the species. Germination phenology is highly correlated with the responses of seeds to the yearly temperature cycle. Species with winter and summer annual life cycles have predictable germination characteristics, but monocarpic and polycarpic perennials do not. Several dormancy types may be found in a given habitat, and narrowly endemic and widely-distributed species in the same genus may have similar germination characteristics. Within some families there is a tendency for a particular type of seed-temperature response to be very important, but frequently this is related to the predominance of a given life cycle type in the family.     

Fire studies in mallee (Eucalyptus spp.) communities of western New South Wales: The effects of fires applied in different seasons on herbage productivity and their implications for management

The post-fire development of herbaceous understoreys of Tnodia/mallee burnt in different seasons was studied over a 3 year period in south-western New South Wales, Australia. Near Pooncarie, NSW, a 3 year sequence of above-average rainfall immediately following burning in either the spring, autumn or winter resulted in a substantial increase in species richness and herbage dry matter production. Plots burnt in the spring produced significantly more dry matter than either the autumn or the winter-burnt plots but only in the first growing season after burning. Some species such as Haloragis odontocarpa behaved as ephemerals, being abundant in the first post-fire season, particularly after spring fire, before becoming inconspicuous. Short-lived grasses, such as Stipa species, then became relatively abundant except during drought. Other experiments confirmed the potential of fire to increase herbage diversity and productivity when there was adequate rainfall. During drought, post-fire herbage production was low and the dominant perennials, viz. mallee Eucalyptus and Triodia irritans, regenerated at the expense of the herbs. There was a strong negative correlation between combined mallee and Triodia cover and total herbage cover over time. Seedling recruitment of Triodia irritans was significantly higher on plots burnt in the spring at Pooncarie.     

Restoring a Mediterranean‐climate shrub community with perennial species reduces future invasion

Successful restoration of an invaded landscape to a diverse, invasion‐resistant native plant community requires determining the optimal native species mix to add to the landscape. We manipulated native seed mix (annuals, perennials, or a combination of the two), while controlling the growth of non‐native species to test the hypothesis that altering native species composition can influence native establishment and subsequent non‐native invasion. Initial survival of native annuals and perennials was higher when seeded in separate mixes than when combined, and competition between the native perennials and annuals led to lower perennial cover in year 2 of mixed‐seeded plots. The plots with the highest perennial cover had the highest resistance to invasion by Brassica nigra. To clarify interactions among different functional groups of natives and B. nigra, we measured competitive interactions in pots. We grew one native annual, one native perennial, and B. nigra alone or with different competitors and measured biomass after 12 weeks. Brassica nigra was the strongest competitor, limiting the growth of all native species, and was not impacted by competition with native annuals or perennial seedlings. Results from the potted plant experiment demonstrated the strong negative influence of B. nigra on native seedlings. Older native perennials were the strongest competitors against invasive species in the field, yet perennial seedling survival was limited by competition with native annuals and B. nigra. Management action that maximizes perennial growth in early years may lead to a relatively more successful restoration and the establishment of an invasion‐resistant community.     

Fate of plants from buried seeds on Volcano Usu,Japan, after the 1977–1978 eruptions

The eruptions of 1977–1978 on Mount Usu in northern Japan resulted in the almost complete destruction of vegetation by a 1–3-m-thick accumulation of volcanic deposits. Erosion created gullies that removed these deposits and frequently exposed the old original soil. I previously confirmed that seedlings of 14 species (eight annual and six perennial herbs) emerged from seeds buried in the original topsoil. To clarify the role of the seed bank on volcanic succession, I monitored seedlings of seed bank species from 1983 to 1992. Nearly all the annuals, such as Polygonum longisetum and Rorippa islandica, originated from the seed bank. However, seed supply from the seed bank declined with time. Because seedling mortality was extremely high and reproductive success was low, due mostly to the instability of ground surface, these annuals emerged for several years but disappeared after 1989. The nitrogen-fixing and stoloniferous perennials, Trifolium repens and Lotus corniculatus var. japonicus, were derived only from the seed bank. They have gradually increased in cover and have become large enough to flower. The other perennials derived from the seed bank, most of which produce short rhizomes, did not increase in cover annually and thus contributed less to revegetation than nitrogen-fixing stoloniferous species. The fate of perennials seems to be dependent on the types of rhizome and stolon systems. Although recruitment from the seed bank was restricted to gullies, the seed bank was the major source of annuals and of nitrogen-fixing perennials in the gullies. Succession in the gullies was substantially different from that which occurred outside the gullies where plants mostly originated from seed immigration and vegetative reproduction from surviving plants, and annual and nitrogen-fixing plants were absent.     

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.     

三峡水库蓄水对消涨带植被的初步影响

三峡库区消涨带植被一直受到广泛关注。为了研究水库蓄水对175 m以下消涨带原有植被的潜在影响,2008年8月在三峡大坝上游长江干流从巴南到秭归12个监测点,设立68个5 m×5 m永久监测样方,比较被水淹过的156 m以下与当时尚未水淹的156～175 m地段植被物种组分、物种多样性和草本层生物量的差异。结果表明：被水淹过与尚未水淹的植被物种组分存在显著差异,DCA可以将二者明显分开,156 m以上尚未水淹的植被以灌丛为主;156 m以下被水淹过的植被以草丛为主,优势种包括多年生草本：狗牙根（Cynodon dactylon）、牛鞭草（Hemarthria altissima）和硬秆子草（Capillipedium assimile）;一年生草本：狗尾草（Setaria viridis）、毛马唐（Digitaria ciliaris）和千金子（Leptochloa chinensis）。木本植物基本死亡,只有枫杨（Pterocarya stenoptera）、牡荆（Vitex negundo）和地瓜（Ficus tikoua）存活。此外,被水淹过的植被中灌木物种数比例显著降低,而一年生草本物种数比例显著增加。被水淹过和尚未水淹植被的物种多样性也存在显著差异,被水淹过植被物种总数、灌木和多年生草本物种数显著低于尚未被水淹过的植被,但一年生草本物种数没有显著差异。被水淹过的植被草本层生物量与尚未水淹的植被没有显著差异。可见,水淹显著改变了消涨带原有植被物种组分和物种多样性,但一年生草本物种数和草本层生物量没有显著差异。未来175 m以下的植被中,草本尤其是一年生草本将占据优势。在消涨带植被恢复选择适应水淹生境物种过程中,应分别从植物的生活史、生理学和形态学等角度进行筛选,尤其应重视生活史适应策略植物的应用。被水淹过的156 m以下现有优势草本和存活的灌木可以作为三峡库区未来消涨带植被恢复的备选物种。     

Differential utilization of summer rains by desert plants

Summary Seasonal changes in the hydrogen isotope ratios of xylem waters were measured to determine water sources used for growth in desert plants of southern Utah. While all species used winter-spring recharge precipitation for spring growth, utilization of summer rains was life-form dependent. Annuals and succulent perennials exhibited a complete dependence on summer precipitation. Herbaceous and woody perennial species simultaneously utilized both summer precipitation and remaining winter-spring precipitation, with herbaceous species much more reliant on the summer precipitation component. Several of the woody perennials exhibited no response to summer precipitation. Currently, precipitation in southern Utah is evenly partitioned between winter and summer time periods; however, global circulation models predict that summer precipitation will increase in response to anticipated climate change. Our data indicate that components within the community will differentially responde to the change in precipitation patterns. These results are discussed in relation to impact on competition and possible changes in community structure.     

Invading Monotypic Stands of Phalaris arundinacea: A Test of Fire, Herbicide, and Woody and Herbaceous Native Plant Groups

Phalaris arundinacea L. is an aggressive species that can dominate wetlands by producing monotypic stands that suppress native vegetation. In this study invasion windows were created for native species in monotypic stands of P. arundinacea with either fire or herbicide. Three native species groups, herbaceous plants, herbaceous seeds, and woody shrubs, were planted into plots burned or treated with herbicide in the early spring. Fire did not create an effective invasion window for native species; there was no difference in P. arundinacea root and shoot biomass or cover between burned and control plots (p≥ 0.998). Herbicide treatment created an invasion window for native species by reducing P. arundinacea root and shoot biomass for two growing seasons, but that invasion window was fast closing by the end of the second growing season because P. arundinacea shoot biomass had nearly reached the shoot biomass levels in the control plots (p= 0.053). Transplant mortality, frost, and animal herbivory prevented the herbaceous species and woody seedlings from becoming fully established in the plots treated with herbicide during the first year of the experiment. Transplanted monocots had a greater survival than dicots. By the second growing season the herbaceous group had the greatest mean areal cover (5%), compared to the woody seedlings (3%) and seed group (0%). Long‐term monitoring of the plots will determine whether the herbaceous transplants will compete effectively with P. arundinacea and whether the woody species will survive, shade the P. arundinacea, and accelerate forest succession.     

Reproductive competence from an annual and a perennial perspective

Plants at early stages of development undergo a juvenile phase during which they are not competent to flower in response to environmental stimuli. The length of this phase varies among species and is extended in perennial plants particularly. In annuals, temporal changes in expression of microR156 (miR156), miR172, and their targets are correlated with the transition from the juvenile to the adult phase and flowering. This developmental transition in perennials is probably more complex than in other plants and the molecular mechanisms are less well understood. In addition, once perennials become adult and capable of reproduction they still keep some meristems in the vegetative state that contribute to their polycarpic growth habit. Juvenility and polycarpy, although considered as two different processes in perennials, might be related.     

Effect of night temperature on daytime stomatal conductance in early and late successional plants

Summary Sixteen annuals, biennials, and herbaceous and woody perennials characteristic of early and late successional old field ecosystems and upland and floodplain habitats were analyzed for their response of stomatal conductance to changes in night temperature. Early successional species that germinate in early spring when temperatures are low, but above freezing are insensitive to cool nights, i.e., their conductance in the following days is unaffected by low night temperature. Later spring and summer-emerging species' stomatal conductance is inhibited by low temperatures. Tree species show the same effects and in some an enhancement of stomatal conductance by low night temperatures was observed. However, adaptive differences in response to night temperatures appear related to both phenology of germination and growth and habitat types.