Article

Evaluating the role of phenology in managing urban invasions: A case study of Broussonetia papyrifera

Authors:
  • Jain Irrigation Systems Ltd., Confedreration of Horticulture Associations of India(CHAI), New Delhi , formerly DDG(Hort), ICAR, vice- chancellor, RAU Pusa,
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Abstract

Multidisciplinary approaches are required for the management of invasive woody species in urban areas. In this context, phenological studies are a useful tool to understand tree development and devise suitable management strategies under urban forestry conditions. The significant role of phenology in attributing competitive advantages to invasive alien species has also been long recognized by community ecologists. Therefore, phenological calendars of invasive species may help in their detection, management and mitigation. In the present study, an attempt has been made to understand the invasive potential of an emerging invasive tree of urban landscapes, Broussonetia papyrifera (paper mulberry; Moraceae), through its phenological assessment by using a standardized numerical scale, BBCH (Biologische Bundesanstalt, Bundessortenamt, CHemische Industrie). The tree is native to southeastern and eastern Asia and the Pacific Islands, and is rapidly spreading across various tropical and subtropical regions. The phenology of B. papyrifera was described in eight principal growth stages (PGSs) with two developmental cycles in a year, presented as primary (January-June) and secondary (June-November) flushes. The observations were further supported by the corresponding dates, photographs, meteorological data (air temperature, precipitation, and photoperiod), and climatic water balance of the study area. Results suggest that the occurrence of two developmental cycles, a prolonged reproductive period, strategy to attract frugivores with brightly colored pulpy fruits, and ability to survive under a wide temperature range help in the spread of B. papyrifera. The phenological scale provided in this study describes accurate and precise developmental stages of the tree that can be used to devise efficient management strategies for its control in urban areas. The information can also be exploited for assessing the climatic conditions required for its prevalence, predicting its future geographic range, and further research.

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Longan (Dimocarpus longan) is an important evergreen fruit crop grown in tropical and subtropical climates, with a clear expanding potential, but with a poorly described phenology. In this work, the different phenological growth stages of longan are characterized according to the BBCH (Biologische Bundesanstalt, Bundessortenamt und Chemische industrie) scale. From vegetative bud dormancy to fruit harvest, seven main growth stages are described for bud, leaf and shoot development, inflorescence emergence, flowering, fruit development and fruit maturity. Within them 41 secondary growth stages are described. The BBCH code will be an important tool to assist the development and implementation of longan management protocols and to standardize observations made in different edaphoclimatic and experimental conditions.
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Broussonetia papyrifera (L.) Vent. (Moraceae), or paper mulberry, is a species of cultural importance in South East Asia, East Asia and the Pacific. Originally from mainland South East Asia or East Asia, this plant was introduced into the Pacific range by prehistoric Austronesian voyagers. We used non-coding internal transcribed spacer (ITS) sequences of nuclear ribosomal DNA and inter-simple sequence repeat (ISSR) on 79 samples of B. papyrifera from different islands of Remote Oceania, and South East Asia and East Asia. Our results show an absence of genetic diversity in the introduced range of Remote Oceania, with the sole exception of Hawaii. By contrast, Asian samples show genetic diversity. The data obtained suggest a prehistoric human-mediated introduction of this species from East Asia to Remote Oceania and a second, possibly historic, human-mediated introduction to Hawaii.
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Phenological documentation and plant invasion are key contemporary issues in the biological sciences. Recognizing this, the present paper describes the phenological stages of an invasive species, Sapium sebiferum (L.) Roxb. A tree of the species was marked, and daily visits were made for recording its phenology. The Biologische Bundesanstalt, Bundessortenamt and CHemical industry (BBCH) scale was used for documenting the same. Of the total ten principal stages, eight were observed in the species. These stages started with bud development (stage 01) in the month of March and ended with leaf fall (stage 97) in the month of December. Descriptions of the phenological stages and their dates are provided in the text. It is hoped that such a phenological scale will be helpful in managing plant invasion.
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Following its introduction in the late 1960s, Broussonetia papyrifera L. Vent. Moraceae (paper mulberry) has emerged as a major exotic invasive species in Ghana's forest ecosystems. This study was carried out to assess the effects of B. papyrifera invasion on community composition in forest and forest-savannah transition ecosystems. Comparative and removal experiments were conducted in paired B. papyrifera invaded versus uninvaded plots. In the comparative assessment, species composition was found to be similar in both invaded and uninvaded plots. However, relative per cent cover of resident species and guilds including Chromolaena odorata, indigenous broadleaves and indigenous grasses were significantly lower in invaded plots. Seven months after B. papyrifera was experimentally removed from invaded stands, cover by indigenous broadleaves increased by 35%, as against only 5% in control plots at the forest site. However, at the transition site, the increase in per cent cover of indigenous broadleaves (18%) was not significantly different from control (2.5%) plots. We conclude that B. papyrifera has the capacity to reduce the abundance of indigenous broadleaf species, although its removal is more likely to favour regeneration in a forest than a forest-savannah transition ecosystem.
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The wide knowledge gaps in invasion biology research that exist in the developing world are crucial impediments to the scientific management and global policymaking on biological invasions. In an effort to fill such knowledge gaps, we present here an inventory of the alien flora of India, based on systematic reviews and rigorous analyses of research studies (ca. 190) published over the last 120 years (1890–2010 AD), and updated with field records of the last two decades. Currently, the inventory comprises of 1,599 species, belonging to 842 genera in 161 families, and constitutes 8.5% of the total Indian vascular flora. The three most species-rich families are Asteraceae (134 spp.), Papilionaceae (114 spp.) and Poaceae (106 spp.), and the three largest genera are Eucalyptus (25 spp.), Ipomoea (22 spp.), and Senna (21 spp.). The majority of these species (812) have no report of escaping from cultivation. Of the remaining subset of 787 species, which have either escaped from intentional cultivation, or spread after unintentional introduction, casuals are represented by 57 spp., casual/naturalised by 114 spp., naturalised by 257 spp., naturalised/invasive by 134 spp., and invasive by 225 spp. Biogeographically, more than one-third (35%) of the alien flora in India has its native ranges in South America, followed by Asia (21%), Africa (20%), Europe (11%), Australia (8%), North America (4%); and cryptogenic (1%). The inventory is expected to serve as the scientific baseline on plant invasions in India, with implications for conservation of global biodiversity.
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ABSTRACT The ecology of seed dispersal by vertebrates has been investigated extensively over recent decades, yet only limited research has been conducted on how suites of invasive plants and frugivorous birds interact. In this review, we examine how plant fruit traits (morphology, colour and display, nutritional quality, accessibility and phenology), avian traits (fruit handling techniques, gut passage time and effect, bird movements and social behaviour and dietary composition) and landscape structure (fruit neighbourhood, habitat loss and fragmentation and perch tree effects) affect frugivory and seed dispersal in invasive plants. This functional approach could be used to develop generic models of seed dispersal distributions for suites of invasive plant species and improve management efficiencies. Four broad research approaches are described that could direct management of bird-dispersed invasive plants at the landscape scale, by manipulating dispersal. First, research is needed to quantify the effect of biological control agents on dispersal, particularly how changes in fruit production and/or quality affect fruit choice by frugivores, dispersal distributions of seed and post-dispersal processes. Second, we explore how seed dispersal could be directed, such as by manipulating perch structures and/or vegetation density to attract frugivorous birds after they have been foraging on invasive plant fruits. Third, the major sources of seed spread could be identified and removed (i.e. targeting core or satellite infestations, particular habitats and creating barrier zones). Fourth, alternative food resources could be provided for frugivores, to replace fruits of invasive plants, and their use quantified.
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ABSTRACT The traditional approach to understanding invasions has focused on properties of the invasive species and of the communities that are invaded. A well-established concept is that communities with higher species diversity should be more resistant to invaders. However, most recently published field data contradict this theory, finding instead that areas with high native plant diversity also have high exotic plant diversity. An alternative environment-based approach to understanding patterns of invasions assumes that native and exotic species respond similarly to environmental conditions, and thus predicts that they should have similar patterns of abundance and diversity. Establishment and growth of native and exotic species are predicted to vary in response to the interaction of plant growth rates with the frequency and intensity of mortality-causing disturbances. This theory distinguishes between the probability of establishment and the probability of dominance, predicting that establishment should be highest under unproductive and undisturbed conditions and also disturbed productive conditions. However, the probability of dominance by exotic species, and thus of potential negative impacts on diversity, is highest under productive conditions. The theory predicts that a change in disturbance regime can have opposite effects in environments with contrasting levels of productivity. Manipulation of productivity and disturbance provides opportunities for resource managers to influence the interactions among species, offering the potential to reduce or eliminate some types of invasive species.
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The spread of many invasive plants is facilitated through seed dispersal by frugivorous animals. The effectiveness of various frugivores as dispersers of the seeds of Melia azedarach, a highly invasive alien tree species, was evaluated in South Africa in savanna and bushveld vegetation. During 264 h of observation, seven bird species and one bat species were recorded foraging on fruiting trees of M. azedarach. The most common visitors were the dark-capped bulbul (Pycnonotus barbatus) followed by Wahlberg's epauletted fruit bat (Epomophorus wahlbergi), but both species droppednearly as many seeds as they dispersed. Knysna turaco (Tauraco corythaix) dispersed the highest number of fruits per minute, but occurred in low abundance in our study sites. Seed germination differed significantly between de-pulped fruits and untreated fruits after 2 mo, but was similar after 4 mo. Germination success did not differ between animal-handled and hand-depulped fruits. In contrast to the high germination success in the greenhouse, seedlings showed very low recruitment in the field. Thus, M. azedarach seems likely to benefit from frugivores (particularly dark-capped bulbul and Wahlberg's epauletted fruit bat) dispersing seeds to suitable microsites.
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1 Species attributes have been used to explain invasion patterns assuming the prevalence of biological mechanisms, although this approach often suffers several methodological and conceptual limitations, such as local idiosyncrasies, differences among habitats, phylogenetic constraints and insufficient sample size. 2 The relative importance of 15 species traits for explaining the abundance over 350 naturalized alien plant species was assessed across five Mediterranean islands (Corsica, Crete, Majorca, Malta and Sardinia). A comparative analysis accounting for phylogeny was used to examine variation in semi-quantitative estimates of species abundance in comparable habitats across the five island floras. 3 Species were divided into those with affinity for semi-natural, agricultural and ruderal habitats. Both vegetative and reproductive attributes were evaluated for individual islands and averaged across all islands. 4 Vegetative propagation, large leaf size, summer flowering, long flowering period and dispersal by wind or vertebrates were positively associated with average alien abundance across all five islands. Fewer significant trends were found in island-specific patterns. 5 The relative importance of a few reproductive traits is reflected in over-representation of Caryophyllales, Asterales and Poales (late flowering, large seed size and anemochory). Although significant covariation in traits was found there was no evidence for welldefined invasive syndromes. 6 Succulence was important in ruderal habitats, long flowering period in agricultural habitats and vertebrate seed dispersal in semi-natural habitats, suggesting that empty niches, avoidance of competitors and exploitation of mutualists, respectively, are important. 7 The study highlights the importance of estimating invasion success across a wide region, but analyses of specific invasion stages are also needed. Reproductive traits, which may be more relevant for long-distance colonization, and vegetative traits, which determine local dominance and persistence, were, nevertheless, both related to abundance within islands.
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Biological invasion caused by the non-indigenous plant species have been considered as one of the major threat to the native vegetation and its diversity at local, regional and global level and its effect on native forest ecosystem is regarded as one of the most critical issues confronting environmental conservationists. This study addresses the influence of Broussonetia papyrifera on the native scrub forest at the Himalayan Foothills, Islamabad, Pakistan and its relationship with ecological gradients which are important in terms of its spatial distribution. Floristic species composition and environmental factors were measured from 77 plots from two sites of the scrub forest at the lower elevation of Margalla Hills National Park Islamabad. Agglomerative hierarchical Cluster Analysis (CA) was used for species assemblage patterns and ordination analyses such as Detrended Correspondence Analysis (DCA) and Canonical Correspondence Analysis (CCA) was used to establish the relationship with the underlying ecological gradients. CA divided the plots into three vegetation zones: a vegetation zone comprised of species of native scrub forest dominated by Acacia modesta; a transition vegetation zone where B. papyrifera was present either in scattered form amongst the scrub forest species or showed patchy distribution and invasive vegetation zone dominated by B. papyrifera. Man Whitney U-test was used to find out if vegetation zones identified by CA could be significantly different from each other based on the measured environmental factors. Factor Analysis (FA)/Principal Components Analysis (PCA) were used to identify set of environmental factors/predictors, which can best discriminate vegetation zones. FA/PCA (raw varimax rotated) on the environmental factors renders three varifactors with eigenvalues higher than 1.0 accounting for over 72% of total variance. Multivariate analyses indicated that the spatial distribution of B. papyrifera is related to edaphic factors such as the soil texture, organic matter and moisture contents. The distribution was insensitive to the topographic factors. The probable consequences of B. papyrifera invasion for future scrub forest composition and plant species diversity are also discussed. The findings of this research will be used in formulating scientifically sound management and conservation strategy for the forest ecosystems of the Margalla Hills National Park Islamabad.
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This study aims to investigate the anaerobic digestion (AD) performance of Broussonetia papyrifera, a wide distributed nitrogen-rich and lignocellulosic-like plant, for methane production. The fermentation was conducted with the volatile solid (VS) concentration at 0.5%, 1.0%,1.5%,2.0% and 4.0% under 37°C and 55°C for 30 days, respectively. Results suggested that the feedstock concentration at 2.0% and fermentation temperature at 55°C was optimal for the AD treatment of Broussonetia papyrifera. A maximal specific methane yield was achieved at 277.72 ± 15 mL/g VS, which was 94.71% higher than that for the maximum specific methane under the thermophile condition, likely because that the C/N was more suitable compared with other lignocellulosic biomass. A kinetic study suggested that the modified Gompertz equation was reliable (determination coefficients R² greater than 0.97) to describe the kinetic behavior of anaerobic digestion of Broussonetia papyrifera. In general, Broussonetia papyrifera was proved to be a suitabe feedstock for anaerobic digestion for methane production.
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Since Schinus molle (Peruvian pepper tree) was brought to Southern California, the dioecious tree has been widely adopted in urban forests and has become invasive. Prior to our study, it was unknown whether S. molle influences plant species in its understory through allelopathy as it does in other geographic areas. The present study sought to determine if S. molle has negative effects on three native and three non-native, invasive understory species. For two separate experiments seeds of six plant species were sowed in mulch (leaf litter) and soil collected from both genders of S. molle. Because a prior study showed that native succulents were most inhibited by males, we expected male mulch and soil would affect native and invasive species more than that of females; however, this prediction was not supported. Male leaf litter of S. molle stimulated the shoot biomass for Amsinckia intermedia, Bromus madritensis, Brassica nigra, and Stipa pulchra. Male soils reduced the shoot emergence of B. madritensis by 34%, however, both genders of soil did not influence the root and shoot biomass of any species. Additionally, field observations in three urban nature reserves recorded 20 native and 10 invasive plant species beneath S. molle canopies. The present study showed S. molle did not have the same influence on species as previously reported for native succulents.
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Soil nitrogen-fixing bacterial communities (SNB) can increase the level of available soil N via biological N-fixation to facilitate successful invasion of several invasive plant species (IPS). Meanwhile, landscape heterogeneity can greatly enhance regional invasibility and increase the chances of successful invasion of IPS. Thus, it is important to understand the soil micro-ecological mechanisms driving the successful invasion of IPS in heterogeneous landscapes. This study performed cross-site comparisons, via metagenomics, to comprehensively analyze the effects of Solidago canadensis invasion on SNB in heterogeneous landscapes in urban ecosystems. Rhizospheric soil samples of S. canadensis were obtained from nine urban ecosystems [Three replicate quadrats (including uninvaded sites and invaded sites) for each type of urban ecosystem]. S. canadensis invasion did not significantly affect soil physicochemical properties, the taxonomic diversity of plant communities, or the diversity and richness of SNB. However, some SNB taxa (i.e., f_Micromonosporaceae, f_Oscillatoriaceae, and f_Bacillaceae) changed significantly with S. canadensis invasion. Thus, S. canadensis invasion may alter the community structure, rather than the diversity and richness of SNB, to facilitate its invasion process. Of the nine urban ecosystems, the diversity and richness of SNB was highest in farmland wasteland. Accordingly, the community invasibility of farmland wasteland may be higher than that of the other types of urban ecosystem. In brief, landscape heterogeneity, rather than S. canadensis invasion, was the strongest controlling factor for the diversity and richness of SNB. One possible reason may be the differences in soil electrical conductivity and the taxonomic diversity of plant communities in the nine urban ecosystems, which can cause notable shifts in the diversity and richness of SNB.
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Paper mulberry is a valuable woody species with a good chilling tolerance. In this study, phosphoproteomic analysis, physiological measurement and mRNA quantification were employed to explore the molecular mechanism of chilling (4 ?C) tolerance in paper mulberry. After chilling for 6 hours, 427 significantly changed phosphoproteins were detected in paper mulberry seedlings without obvious physiological injury. When obvious physiological injury occurred after chilling for 48 hours, a total of 611 phosphoproteins were found to be significantly changed at the phosphorylation level. Several protein kinases, especially CKII, were possibly responsible for these changes, according to conserved sequence analysis. The results of GO analysis showed that phosphoproteins were mainly responsible for signal transduction, protein modification and translation during chilling. Additionally, transport and cellular component organization were enriched after chilling for 6 and 48 hours, respectively. Based on the protein-protein interaction network analysis, a protein kinases and phosphatases hub protein (P1959) was found to be involved in cross-talk between Ca2+, BR, ABA and ethylene-mediated signaling pathways. We also highlighted the phosphorylation of BpSIZ1 and BpICE1 possibly impacted on the CBF/DREB-responsive pathway. From these results, we developed a schematic for the chilling tolerance mechanism at phosphorylation level.
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Tree phenology describes complex tree growth and developmental processes. Phenological scales are a useful tool for understanding tree growth/establishment and to evaluate tree phenology management practices under urban forestry conditions. Scales such as (1) the BBCH scale (Biologische Bundesantalt, Bundessortenamt und Chemische Industrie) and a (2) landmark scale (developed by the authors) were used to characterize the phenological changes occurring in the capuli cherry Prunus serotina subsp. capuli (Cav.) McVaugh growing under tropical conditions in Bogotá, Colombia. The BBCH scale is an applicable tool that describes whole tree canopy phenological events, and the landmark scale is a useful descriptor of individual asynchronous stems, which occur as ephemeral flushes at different portions of the canopy. Our results depicted several principal advantages to using the BBCH scale. The landmark scale described fewer vegetative, reproductive, and fruiting stages when compared to the BBCH scale. The landmark scale included the timing of each stage/event and the BBCH failed to do so. Thus, both scales differ in number of stages, coding system, timing of each individual stage, and synchronous (BBCH scale) vs asynchronous (Landmark scale) events. Both scales are applicable, useful and meaningful to the study site and the field of urban forestry. This study helps to expand our knowledge of the capuli cherry's phenology and urban forestry applications.
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The accurate prediction of winter injury caused by low-temperature events is a key component of the effective cultivation of woody and herbaceous perennial plants. A common method employed to visualize geographic patterns in the severity of low-temperature events is to map a climatological variable that closely correlates with plant survival. The U.S. Department of Agriculture Plant Hardiness Zone Map (PHZM) is constructed for that purpose. We present a short history of PHZM development, culminating in the recent production of a new, highresolution version of the PHZM, and discuss how such maps relate to winterhardiness per se and to other climatic factors that affect hardiness. The new PHZM is based on extreme minimum-temperature data logged annually from 1976 to 2005 at 7983 weather stations in the United States, Puerto Rico, and adjacent regions in Canada and Mexico. The PHZM is accessible via an interactive website, which facilitates a wide range of horticultural applications. For example, we highlight how the PHZM can be used as a tool for site evaluation for vineyards in the Pacific northwestern United States and as a data layer in conjunction with moisture-balance data to predict the survival of Yugoslavian woody plants in South Dakota. In addition, the new map includes a zip code finder, and we describe how it may be used by governmental agencies for risk management and development of recommended plant lists, by horticultural firms to schedule plant shipments, and by other commercial interests that market products seasonally.
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Phenology, the timing of annually recurrent reproductive biological events, provides a critical signal of climate variability and change effects on plants. Considerable work over the past five decades has quantified the extent to which plant phenophases are responding to local changes in temperature and rainfall. Originally undertaken through the analysis of ground-based phenological observations, the discipline has more recently included phenophase indicators from satellite images and digital repeat photography. With research advances it has become evident that the responses of plant phenology to climate variability and change are both location- and species-specific. The extent to which plants are affected by changes in temperature and rainfall, their intrinsic adaptation capacity, will ultimately determine the potential for sustained ecological stability and food security. We review methodological approaches to plant phenological-climate change over time, analyse the regions and phenophases for which climate variability demonstrates a clear causal role, and finally reflect on the applications of phenological climate change investigations in broader biogeographical contexts.
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A study was carried out in Panama on Ficus insipida and F. yoponensis (subgenus Pharmacosycea), F. obtusifolia and F. costaricana (subgenus Urostigma), Cecropia insignis and Poulsenia armata. Overall, phenological activities were more pronounced in the drier half of the year, but at least some leaf shed, leaf flush and fruit production took place throughout the year in most species. As a family, as species and, frequently, as individuals, these Moraceae exhibited phenological traits that would tend to make their new leaves and/or fruits available to primary consumers in this forest for much or all of an annual cycle. The reproductive phenology of Ficus species in this study contrasted with that of C. insignis and P. armata in that individuals of the Ficus species could produce massive fruit crops at any time of year. This temporal flexibility with respect to the timing of fruit crops appears to relate, at least in part, to the obligate mutualism of fig trees with wasp pollinators. -from Author
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To elucidate climate-driven changes in leaf-out phenology and their implications for species invasions, we observed and experimentally manipulated leaf out of invasive and native woody plants in Concord, MA, USA. Using observations collected by Henry David Thoreau (1852-1860) and our own observations (2009-2013), we analyzed changes in leaf-out timing and sensitivity to temperature for 43 woody plant species. We experimentally tested winter chilling requirements of 50 species by exposing cut branches to warm indoor temperatures (22°C) during the winter and spring of 2013. Woody species are now leafing out an average of 18 d earlier than they did in the 1850s, and are advancing at a rate of 5 ± 1 d °C(-1) . Functional groups differ significantly in the duration of chilling they require to leaf out: invasive shrubs generally have weaker chilling requirements than native shrubs and leaf out faster in the laboratory and earlier in the field; native trees have the strongest chilling requirements. Our results suggest that invasive shrub species will continue to have a competitive advantage as the climate warms, because native plants are slower to respond to warming spring temperatures and, in the future, may not meet their chilling requirements.
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Vegetation develops in response to many different stimuli including climatic, edaphic, and cultural conditions. The degree to which the distribution of vegetation can be explained on the basis of climatic conditions depends in part on the proper selection of active climatic factors. Temperature and precipitation by themselves are poor descriptors of climate. Potential evapotranspiration, water surplus, and water deficit (combined in a moisture index) are found to be more useful. An analysis of the vegetation distribution in conterminous United States shows that the Thornthwaite moisture index and annual potential evapotranspiration are not only able to differentiate among the mid-latitude grassland, forest, and desert biochores but also to provide climatic limits for formation classes within each of the biochores. For example, it is possible to identify on the basis of numerical data an oak-hickory climate as distinct from the birch-maple, and a bunch grass climate distinct from the sagebrush. The problem of non-climatically controlled vegetation associations such as the low latitude pines and savanna are also discussed.
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Bitou bush, Chrysanthemoides monilifera (L.) Norlindh ssp. rotundata (DC.) Norlindh, is an invasive weed of coastal vegetation types in eastern Australia. It produces fleshy fruits, which are dispersed by birds and mammals. Compared with a range of vertebrate-dispersed fruits of native plant species that co-occur with naturalised C. monilifera, or occur elsewhere in south-eastern Australia, C. monilifera fruits and endocarps are intermediate in all dimensions and in the ratio of endocarp to pulp. C. monilifera fruits contain moderate quantities of sugars, but low quantities of protein and fat relative to many native fruits. Phenology, however, substantially distinguishes C. monilifera from co-occurring native plant species. C. monilifera produces fruits throughout the year, with the peak of production occurring in autumn to winter, a time when few native species in coastal vegetation produce fruits. Additionally, 9 of the 12 native species that do fruit, at least partly, over this period, differ substantially from C. monilifera in fruit morphology. C. monilifera, along with other invasive species, has substantially altered the temporal pattern of fruit availability in coastal vegetation. These fruiting characteristics have rendered C. monilifera fruits attractive to frugivores and have contributed to its successful spread.
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six phenological patterns are recognized in the flora of a tropical savanna in the Venezuelan Llanos. The dominant perennial grasses and sedges have an alternation between a period of intensive tiller production, shoot growth and sexual reproduction during the rainy season, and a phase of semirest during the dry season, when shoots decline and tiller development decreases to a minimum. An opposite strategy is displayed by the tree species; they have a phase of full leaf development and maximum assimilation during the rainy season, but without any shoot growth or leaf formation, while in the dry season they renew the foliage and flower. Annuals constitute a numerically-significant part of the savanna flora, and have been ascribed to two different phenological groups. The long-cycle annuals form one group and these have an active life cycle of 6 to 7 months, coincident with the rainy season; the other group is formed by the short-cycle annuals or ephemerals, which complete their cycle in only 2 to 4 months during the latter part of the rainy season. Another phenological strategy is displayed by those species which enter into a dormancy phase during drought, when all their above-ground organs dry out; their active phenophases occur during the rainy period. The sixth phenological group is formed by the evergrowing the everflowering plants, i.e. those savanna species without a manifest seasonality. In the semi-deciduous tropical forest which occurs in the same area, two phenological strategies were discerned. The first is displayed by deciduous trees which stay leafless for a part of or for the entire dry season, and which put out a new crop of leaves either just before the onset of rains, or at the beginning or the rainy season. Different reproductive patterns are found in this group, with various species flowering at different times, either during the wet or the dry season. The second strategy in this forest is comprised of evergreen and brevidecidous species which renew their foliage during drought, when both opposite phenophases (leaf flush and leaf drop) proceed simultaneously. Flowering also takes place during the dry season, but only in the brevideciduous species is it directly related to leaf renewal. Considering the total flora of both communities, a major peak of flowering in the middle of the rainy season characterizes the savanna, while the forest has two equally important flowering peaks, one in each season. A discussion of the possible role of various environmental impulses in plant phenology in this region is attempted, considering all seasonal factors, such as daylength, rainfall seasonality, thermoperiodism, extreme temperatures, insolation and radiation. Finally, the conditions favouring the existence and success of each phenological strategy are considered, together with a critical examination of the seasonal characteristics of the two econsystem.
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Information on the geographical distribution of plant invasion has been recorded in detail in some areas of the world; however, in large regions such as South America there are a few, if any, records of the spread of alien plants and invasive species and even less information about their effects on ecosystems at different levels of organization. This study examines the extent to which woody species introduced during the last centuries are invading the Rolling Pampa (which is typical of the entire region of the Argentina pampas) and discusses whether this invasion is related to the species’ genetics or to environmental factors. All woody species were surveyed along landscape corridors (highways and intersecting secondary dirt roads and streams), as well as in farmed fields under three different tillage systems: zero tillage in the entire field for all crops in the rotation (where tillage was replaced by a presowing herbicide application), zero tillage for selected crops, and conventional tillage. Landscape corridors along the roads had been invaded by 40 woody species (mostly trees). On the farmed land, fields under the zero tillage farming system were invaded by seven woody species (three tree species and four shrubs). With zero tillage for select crops only, woody species richness was reduced to three (one tree and two shrub species). In the conventional tillage, there were only three invading species, all shrubs. In both the roadside and riparian corridors, the species with the highest constancy values were Gleditsia triacanthos L., Morus alba L., and Melia azedarach L. In both types of zero tillage farmed fields, M. alba was absent, but G. triacanthos and M. azedarach remained the species with the highest constancy values. Both genetic and ecological factors were important determinants for the invasion of the pampas by woody species. The woody invasion process has reached a point at which the pampean grasslands on the better-drained soils will no longer be restored to a grassland biome without human intervention.
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1. Exotic annual plants are an increasingly important ecological issue and new, creative approaches to management are required. In desert ecosystems of the southwestern USA, the forbs Brassica tournefortii, Erodium cicutarium and Schismus spp. dominate and alter native annual communities. Hand weeding B. tournefortii is currently the most common control method employed, but weeding is inadequate and expensive for managing large-scale invasions. New methods must be developed to conserve and restore desert ecosystems. 2. Exotic annuals in desert systems have rapid germination and phenology compared to natives, indicating that a window for selective control of exotic annuals may occur immediately after exotic seedling emergence. We tested the role of timing in control methods by comparing a cotyledon-stage glyphosate application to a bolting-stage application and to hand weeding B. tournefortii, plus an untreated check. Treatments were tested at two sites dominated by either exotic or native annuals and followed for 2 years; early application was repeated the second year. Cover and richness were evaluated during seedling and peak flowering stages underneath and between shrubs. 3. Early glyphosate application did not affect native cover, but did reduce exotic cover. Late herbicide negatively impacted both exotics and natives. Natives had little positive response, and then only through hand weeding under shrubs, but the same treatment caused an increase in the exotic E. cicutarium. 4.Synthesis and applications. The rapid phenology of exotic annuals may be exploited to control exotics while minimizing impacts on native plants in desert communities. This approach may be useful for other invasions in other ecosystems by species with rapid, early germination, or may act as a supplement to improve the efficacy of existing management regimes.
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Aim  We aim to report what hyperspectral remote sensing can offer for invasion ecologists and review recent progress made in plant invasion research using hyperspectral remote sensing.Location  United States.Methods  We review the utility of hyperspectral remote sensing for detecting, mapping and predicting the spatial spread of invasive species. We cover a range of topics including the trade-off between spatial and spectral resolutions and classification accuracy, the benefits of using time series to incorporate phenology in mapping species distribution, the potential of biochemical and physiological properties in hyperspectral spectral reflectance for tracking ecosystem changes caused by invasions, and the capacity of hyperspectral data as a valuable input for quantitative models developed for assessing the future spread of invasive species.Results  Hyperspectral remote sensing holds great promise for invasion research. Spectral information provided by hyperspectral sensors can detect invaders at the species level across a range of community and ecosystem types. Furthermore, hyperspectral data can be used to assess habitat suitability and model the future spread of invasive species, thus providing timely information for invasion risk analysis.Main conclusions  Our review suggests that hyperspectral remote sensing can effectively provide a baseline of invasive species distributions for future monitoring and control efforts. Furthermore, information on the spatial distribution of invasive species can help land managers to make long-term constructive conservation plans for protecting and maintaining natural ecosystems.