Utilization of invasive tamarisk by salt marsh consumers

Plant invasions of coastal wetlands are rapidly changing the structure and function of these systems globally. Alteration of litter dynamics represents one of the fundamental impacts of an invasive plant on salt marsh ecosystems. Tamarisk species (Tamarix spp.), which extensively invade terrestrial and riparian habitats, have been demonstrated to enter food webs in these ecosystems. However, the trophic impacts of the relatively new invasion of tamarisk into marine ecosystem have not been assessed. We evaluated the trophic consequences of invasion by tamarisk for detrital food chains in the Tijuana River National Estuarine Research Reserve salt marsh using litter dynamics techniques and stable isotope enrichment experiments. The observations of a short residence time for tamarisk combined with relatively low C:N values indicate that tamarisk is a relatively available and labile food source. With an isotopic (¹⁵N) enrichment of tamarisk, we demonstrated that numerous macroinvertebrate taxonomic and trophic groups, both within and on the sediment, utilized ¹⁵N derived from labeled tamarisk detritus. Infaunal invertebrate species that took up no or limited ¹⁵N from labeled tamarisk (A. californica, enchytraeid oligochaetes, coleoptera larvae) occurred in lower abundance in the tamarisk-invaded environment. In contrast, species that utilized significant ¹⁵N from the labeled tamarisk, such as psychodid insects, an exotic amphipod, and an oniscid isopod, either did not change or occurred in higher abundance. Our research supports the hypothesis that invasive species can alter the trophic structure of an environment through addition of detritus and can also potentially impact higher trophic levels by shifting dominance within the invertebrate community to species not widely consumed. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Environmental tolerance of an invasive riparian tree and its potential for continued spread in the southwestern US

Questions: Exotic plant invasion may be aided by facilitation and broad tolerance of environmental conditions, yet these processes are poorly understood in species‐rich ecosystems such as riparian zones. In the southwestern United States (US) two plant species have invaded riparian zones: tamarisk (Tamarix ramosissima, T. chinensis, and their hybrids) and Russian olive (Elaeagnus angustifolia). We addressed the following questions: (1) is Russian olive able to tolerate drier and shadier conditions than cottonwood and tamarisk? (2) Can tamarisk and cottonwood facilitate Russian olive invasion? Location: Arid riparian zones, southwestern US. Methods: We analyzed riparian tree seedling requirements in a controlled experiment, performed empirical field studies, and analyzed stable oxygen isotopes to determine the water sources used by Russian olive. Results: Russian olive survival was significantly higher in dense shade and low moisture conditions than tamarisk and cottonwood. Field observations indicated Russian olive established where flooding cannot occur, and under dense canopies of tamarisk, cottonwood, and Russian olive. Tamarisk and native riparian plant species seedlings cannot establish in these dry, shaded habitats. Russian olive can rely on upper soil water until 15 years of age, before utilizing groundwater. Conclusions: We demonstrate that even though there is little evidence of facilitation by cottonwood and tamarisk, Russian olive is able to tolerate dense shade and low moisture conditions better than tamarisk and cottonwood. There is great potential for continued spread of Russian olive throughout the southwestern US because large areas of suitable habitat exist that are not yet inhabited by this species.     

Rethinking Avian Response to Tamarix on the Lower Colorado River: A Threshold Hypothesis

Many of the world’s large river systems have been greatly altered in the past century due to river regulation, agriculture, and invasion of introduced Tamarix spp. (saltcedar, tamarisk). These riverine ecosystems are known to provide important habitat for avian communities, but information on responses of birds to differing levels of Tamarix is not known. Past research on birds along the Colorado River has shown that avian abundance in general is greater in native than in non‐native habitat. In this article, we address habitat restoration on the lower Colorado River by comparing abundance and diversity of avian communities at a matrix of different amounts of native and non‐native habitats at National Wildlife Refuges in Arizona. Two major patterns emerged from this study: (1) Not all bird species responded to Tamarix in a similar fashion, and for many bird species, abundance was highest at intermediate Tamarix levels (40–60%), suggesting a response threshold. (2) In Tamarix‐dominated habitats, the greatest increase in bird abundance occurred when small amounts of native vegetation were present as a component of that habitat. In fact, Tamarix was the best vegetation predictor of avian abundance when compared to vegetation density and canopy cover. Our results suggest that to positively benefit avian abundance and diversity, one cost‐effective way to rehabilitate larger monoculture Tamarix stands would be to add relatively low levels of native vegetation (～20–40%) within homogenous Tamarix habitat. In addition, this could be much more cost effective and feasible than attempting to replace all Tamarix with native vegetation.     

Insect biological control accelerates leaf litter decomposition and alters short‐term nutrient dynamics in a Tamarix‐invaded riparian ecosystem

Insect herbivory can strongly influence ecosystem nutrient dynamics, yet the indirect effects of herbivore‐altered litter quality on subsequent decomposition remain poorly understood. The northern tamarisk beetle Diorhabda carinulata was released across several western states as a biological control agent to reduce the extent of the invasive tree Tamarix spp. in highly‐valued riparian ecosystems; however, very little is currently known about the effects of this biocontrol effort on ecosystem nutrient cycling. In this study, we examined alterations to nutrient dynamics resulting from beetle herbivory in a Tamarix‐invaded riparian ecosystem in the Great Basin Desert in northern Nevada, USA, by measuring changes in litter quality and decomposition, as well as changes in litter quantity. Generally, herbivory resulted in improved leaf litter chemical quality, including significantly increased nitrogen (N) and phosphorus (P) concentrations and decreased carbon (C) to nitrogen (C:N), C:P, N:P, and lignin:N ratios. Beetle‐affected litter decomposed 23% faster than control litter, and released 16% more N and 60% more P during six months of decomposition, as compared to control litter. Both litter types showed a net release of N and P during decomposition. In addition, herbivory resulted in significant increases in annual rates of total aboveground litter and leaf litter production of 82% and 71%, respectively, under the Tamarix canopy. Our finding that increased rates of N and P release linked with an increased rate of mass loss during decomposition resulting from herbivore‐induced increases in litter quality provides new support to the nutrient acceleration hypothesis. Moreover, results of this study demonstrate that the introduction of the northern tamarisk beetle as biological control to a Tamarix‐invaded riparian ecosystem has lead to short‐term stimulation of nutrient cycling. Alterations to nutrient dynamics could have implications for future plant community composition, and thus the potential for restoration of Tamarix‐invaded ecosystems.     

Northern tamarisk beetle (Diorhabda carinulata) and tamarisk (Tamarix spp.) interactions in the Colorado River basin

Northern tamarisk beetles (Diorhabda carinulata) were released in the Upper Colorado River Basin in the United States in 2004–2007 to defoliate introduced tamarisk shrubs (Tamarix spp.) in the region's riparian zones. The primary purpose was to control the invasive shrub and reduce evapotranspiration (ET) by tamarisk in an attempt to increase stream flows. We evaluated beetle–tamarisk interactions with MODIS and Landsat imagery on 13 river systems, with vegetation indices used as indicators of the extent of defoliation and ET. Beetles are widespread and exhibit a pattern of colonize–defoliate–emigrate, so that riparian zones contain a mosaic of completely defoliated, partially defoliated, and refoliated tamarisk stands. Based on satellite data and ET algorithms, mean ET before beetle release (2000–2006) was 416 mm/year compared to postrelease (2007–2015) ET of 355 mm/year (p < 0.05) for a net reduction of 61 mm/year. This is lower than initial literature projections that ET would be reduced by 300–460 mm/year. Reasons for the lower‐than‐expected ET reductions are because baseline ET rates are lower than initially projected, and percentage ET reduction is low because tamarisk stands tend to regrow new leaves after defoliation and other plants help maintain canopy cover. Overall reductions in tamarisk green foliage during the study are 21%. However, ET in the Upper Basin has shown a steady decline since 2007 and equilibrium has not yet been reached. Defoliation is now proceeding from the Upper Basin into the Lower Basin at a rate of 40 km/year, much faster than initially projected.     

Similar dietary but different numerical responses to nonnative tamarisk (<Emphasis Type="Italic">Tamarix</Emphasis> spp.) by two native warblers

Native species can have a range of responses to nonnative introductions, from negative to positive, and understanding how and why native species respond differently to nonnatives remains an important management challenge. Based on differences and similarities in ecology and behavior, we predicted how abundance and diet of two native warblers, Lucy’s warbler (Oreothlypis luciae) and yellow warbler (Setophaga petechia), would differ in habitats with different amounts of nonnative tamarisk trees and the three nonnative insects obligately dependent on tamarisk (Tamarix spp.). Specifically, we predicted that Lucy’s warblers would have similar densities across sites, yellow warbler densities would be inversely related to tamarisk cover, and both warblers, being generalist insectivores, would incorporate tamarisk biocontrol insects in their diet. Based on point counts and fecal samples at six sites along the Virgin River in the southwestern United States, we found that yellow warblers decreased in abundance with increasing tamarisk cover, while Lucy’s warbler abundance did not and that diet of the two warblers did not differ, with both species exhibiting strong selection for the nonnative tamarisk weevil (Coniatus splendidulus) and weak to no selection for the nonnative tamarisk leafhopper (Opsius stactogalus). Both warblers showed negative selection for the tamarisk beetle (Diorhabda carinulata) and its larvae, even when those insects were 10–100 times more abundant during outbreaks. Although both warblers exploited the novel food resources offered by tamarisk, with those insects contributing half or more of total prey biomass, Lucy’s warblers were better able to maintain densities in tamarisk habitats. We hypothesize this was due to the Lucy’s warbler’s ability to exploit a broader array of habitats surrounding tamarisk sites and its cavity nesting habit that buffers its nests from the higher temperatures and lower humidity of tamarisk-dominated habitat. Our results suggest that predictions based on detailed knowledge of the form and function of native and nonnative species can be used to predict native bird response to nonnatives.     

Rhombomys opimus contribution to the “fertile island” effect of tamarisk mounds in Junggar Basin

Fertile islands are created and maintained by a combination of physical and biologically mediated processes. Plants have been shown to be very important in the formation of fertile islands, and recent research indicates that the activities of burrowing animals have a significant influence on the physiochemical properties of soil that can promote the development of fertile islands. In this paper, we chose tamarisk (Tamarix spp.)??a constructive species that grows in the oasis-desert ecotone, and the great gerbil (Rhombomys opimus)??a widely distributed rodent in Central Asia to study the influence of great gerbils on soil nutrient dynamics in tamarisk mounds. Results indicate that fertile islands exist in tamarisk mounds without burrows of the great gerbil. However, the great gerbil??s burrowing activities promote the fertile island effect in tamarisk mounds: soil nutrients under shrubs with great gerbil activity were significantly higher than inter-mound areas in both surface soil and deep soil from 15 to 50?cm. Available nitrogen in mounds with rodent burrows was over twice as high as in tamarisk mounds without burrows at the same depth. The great gerbil??s burrowing activities promote the concentration of soil nutrients in the tamarisk mounds.     

Diversity and functional responses of nitrogen-fixing microbes to three wetland invasions

Impacts of invasive species on microbial components of wetland ecosystems can reveal insights regarding functional consequences of biological invasions. Nitrogen fixation (acetylene reduction) rates and diversity of nitrogen fixers, determined by genetic fingerprinting (T-RFLP) of the nifH gene, were compared between native and invaded sediments in three systems. Variable responses of nitrogen fixing microbes to invasion by a non-native mussel, Musculista senhousia, and mangrove, Avicennia marina, in Kendall Frost-Northern Wildlife Preserve (Mission Bay) and salt cedar, Tamarisk (Tamarix spp.) in Tijuana Estuary suggest microbes respond to both species- and site-specific influences. Structurally similar invaders (the mangrove and salt cedar) produced different effects on activity and diversity of nitrogen fixers, reflecting distinct environmental contexts. Despite relative robustness of microbial community composition, subtle differences in total diversity or activity of nitrogen fixers reveal that microbes are not immune to impacts of biological invasions, and that functional redundancy of microbial diversity is limited, with significant consequences for functional dynamics of wetlands.     

Riparian bird density decline in response to biocontrol of <Emphasis Type="Italic">Tamarix</Emphasis> from riparian ecosystems along the Dolores River in SW Colorado,USA

Biocontrol of invasive tamarisk (Tamarix spp.) in the arid Southwest using the introduced tamarisk beetle (Diorhabda elongata) has been hypothesized to negatively affect some breeding bird species, but no studies to date have documented the effects of beetle-induced defoliation on riparian bird abundance. We assessed the effects of tamarisk defoliation by monitoring defoliation rates, changes in vegetation composition, and changes in density of six obligate riparian breeding bird species at two sites along the Dolores River in Colorado following the arrival of tamarisk beetles. We conducted bird point counts from 2010 to 2014 and modeled bird density as a function of native vegetation density and extent of defoliation using hierarchical distance sampling. Maximum annual defoliation decreased throughout the study period, peaking at 32–37% in 2009–2010 and dropping to 0.5–15% from 2011–2014. Stem density of both tamarisk and native plants declined throughout the study period until 2014. Density of all bird species declined throughout most of the study, with Song Sparrow disappearing from the study sites after 2011. Blue Grosbeak, Yellow-breasted Chat, and Yellow Warbler densities were negatively related to defoliation in the previous year, while Lazuli Bunting exhibited a positive relationship with defoliation. These findings corroborate earlier predictions of species expected to be sensitive to defoliation as a result of nest site selection. Tamarisk defoliation thus had short-term negative impacts on riparian bird species; active restoration may be needed to encourage the regrowth of native riparian vegetation, which in the longer-term may result in increased riparian bird density.     

The influence of floods and precipitation on <Emphasis Type="Italic">Tamarix</Emphasis> establishment in Grand Canyon,Arizona: consequences for flow regime restoration

Decoupling of climate and hydrology combined with introduction of non-native species creates novel abiotic and biotic conditions along highly regulated rivers. Tamarix, a non-native shrub, dominates riparian assemblages along many waterways in the American Southwest, including the Colorado River through Grand Canyon. We conducted a tree-ring study to determine the relative influences of climate and hydrology on Tamarix establishment in Grand Canyon. Riparian vegetation was sparse and annually scoured by large floods until completion of Glen Canyon Dam, which allowed pioneer species, including Tamarix, to expand. Post-dam floods in the mid-1980s were associated with high Tamarix mortality but also initiated a large establishment event. Subsequent establishment has been low but continuous with some exceptions. From 1984 to 2006 establishment increased during years of high, late-summer flows followed by years of low precipitation. This combination provided moist surfaces for Tamarix establishment and may have caused reduced erosion of seedlings or reduced competition from native plants. Attempts to mimic pre-dam floods for ecosystem restoration through planned flood releases also have affected Tamarix establishment. Early (March 1996) and late (November 2004) restoration floods limited establishment, but a small restoration flood in May 2000 followed by steady summer flows permitted widespread establishment. Flood restoration is not expected to prevent Tamarix spread in this system because historic flood timing in May–July coincides with seed release. To decrease future Tamarix establishment, river managers should avoid floods during peak Tamarix seed release, which encompasses the historic spring and early summer flooding period. Tamarix dominance may be reduced by early spring floods that initiate asexual reproduction of clonal shrubs (e.g., Salix exigua, Pluchea sericea).     

Salt glands of recretohalophyte Tamarix under salinity: Their evolution and adaptation

Here, we studied the evolution of salt glands in 11 species of Tamarix and determined their role in adaptation to saline environments by measuring the effect of NaCl on plant growth and salt gland characteristics. Cluster analysis divided Tamarix species into three types (types I–III) according to salt‐gland characteristics. A phylogenetic tree based on ITS sequences indicated an evolutionary relationship consistent with the geographical distribution of Tamarix. We measured growth under different NaCl conditions (0, 100, 200, and 300 mM) for 40 days in three species (T. gallica, T. ramosissima, and T. laxa) representing the three Tamarix types. With increasing NaCl concentration, the biomass of all species was significantly reduced, especially that of T. gallica. Salt secretion ability and salt‐gland density showed similar trends in three types. The order of salt tolerance was type I > type II > type III. We conclude that during Tamarix adaptation to salinity, salt‐gland evolution followed two directions: one increasing salt‐gland density, and the other increasing salt secretion rate per salt‐gland. This study provides a basis for potential mechanisms of recretohalophyte adaptation to salinity.     

Do riparian plant community characteristics differ between <Emphasis Type="Italic">Tamarix</Emphasis> (L.) invaded and non-invaded sites on the upper Verde River,Arizona?

Invasion by Tamarix (L.) can severely alter riparian areas of the western U.S., which are globally rare ecosystems. The upper Verde River, Arizona, is a relatively free-flowing river and has abundant native riparian vegetation. Tamarix is present on the upper Verde but is a minor component of the vegetation (8% of stems). This study sought to determine whether riparian vegetation characteristics differed between sites where Tamarix was present and sites where Tamarix was absent during the invasion of the upper Verde. We hypothesized that herbaceous understory and woody plant communities would differ between Tamarix present and absent sites. Our hypothesis was generally confirmed, the two types of sites were different. Tamarix present sites had greater abundance of all vegetation, native understory species, graminoids, and native trees, and a positive association with perennial native wetland plant species. Tamarix absent sites had greater abundance of exotic plants and upland adapted plants and an association with greater abiotic cover and litter. These results are contrary to other reports of Tamarix association with depauperate riparian plant communities, and suggest that Tamarix invasion of a watershed with a relatively natural flow regime and a robust native plant community follows similar establishment patterns as the native riparian plant community.     

Selection for genetics‐based architecture traits in a native cottonwood negatively affects invasive tamarisk in a restoration field trial

Climate change and competition from invasive species remain two important challenges in restoration. We examined the hypothesis that non‐native tamarisk (Tamarix spp.) reestablishment after aboveground removal is affected by genetics‐based architecture of native Fremont cottonwood (Populus fremontii) used in restoration. As cottonwood architecture (height, canopy width, number of stems, and trunk diameter) is, in part, determined by genetics, we predicted that trees from different provenances would exhibit different architecture, and mean annual maximum temperature transfer distance from the provenances would interact with the architecture to affect tamarisk. In a common garden in Chevelon, AZ, U.S.A. (elevation 1,496 m), with cottonwoods from provenances spanning its elevation distribution, we measured the performance of both cottonwoods and tamarisk. Several key findings emerged. On average, cottonwoods from higher elevations were (1) two times taller and wider, covered approximately 3.5 times more basal area, and were less shrubby in appearance, by exhibiting four times fewer number of stems than cottonwoods from lower elevations; (2) had 50% fewer tamarisk growing underneath, which were two times shorter and covered 6.5 times less basal area than tamarisk growing underneath cottonwoods of smaller stature; and (3) the number of cottonwood stems did not affect tamarisk growth, possibly because the negative relationship between cottonwood stems and basal area. In combination, these findings argue that cottonwood architecture is affected by local conditions that interact with genetics‐based architecture. These interactions can negatively affect the growth of reinvading tamarisk and enhance restoration success. Our study emphasizes the importance of incorporating genetic and environmental interactions of plants used in restoration.     

The Monstering of Tamarisk: How Scientists made a Plant into a Problem

Dispersal of biota by humans is a hallmark of civilization, but the results are often unforeseen and sometimes costly. Like kudzu vine in the American South, some examples become the stuff of regional folklore. In recent decades, “invasion biology,” conservation-motivated scientists and their allies have focused largely on the most negative outcomes and often promoted the perception that introduced species are monsters. However, cases of monstering by scientists preceded the rise of popular environmentalism. The story of tamarisk (Tamarix spp.), flowering trees and shrubs imported to New England sometime before 1818, provides an example of scientific “monstering” and shows how slaying the monster, rather than allaying its impacts, became a goal in itself. Tamarisks’ drought and salt tolerance suggested usefulness for both coastal and inland erosion control, and politicians as well as academic and agency scientists promoted planting them in the southern Great Plains and Southwest. But when erosion control efforts in Arizona, New Mexico and Texas became entangled with water shortages, economic development during the Depression and copper mining for national defense during World War Two, federal hydrologists moved quickly to recast tamarisks as water-wasting foreign monsters. Demonstrating significant water salvage was difficult and became subsidiary to focusing on ways to eradicate the plants, and a federal interagency effort devoted specifically to the latter purpose was organized and continued until it, in turn, conflicted with regional environmental concerns in the late 1960s.     

Invasive <Emphasis Type="Italic">Tamarix</Emphasis> (Tamaricaceae) in South Africa: current research and the potential for biological control

Most species of Tamarix originate in Eurasia and at least five species have become invasive around the world, including South Africa. However, T. usneoides is indigenous to southern Africa, where the potential for biological control of the invasive species is being investigated. Recent research on the invasive species is reviewed here with particular reference to these South African biocontrol efforts. The successful biological control programme against invasive Tamarix in the USA, using several species of “Tamarisk beetle”, is being used as a guide for the South African research. The South African programme is complicated by firstly, the presence of the indigenous T. usneoides which raises the precision of host-specificity required, and secondly, the introduced and indigenous Tamarix have a high intrinsic value for phytoremediation of mine tailings dams in South Africa. The phylogenetic proximity of these Tamarix species to each other has contributed to this challenge, which has nevertheless been successfully addressed by molecular techniques used to separate the species. In addition, classical morphological techniques have been used to separate the Tamarisk beetles, so that now they can generally be matched to Tamarix tree species. Overall, it is concluded that given the broad knowledge now available on the ecology and identity of both the trees and their biocontrol agents, the prospects for successful biological control of Tamarix in South Africa are good.     

<Emphasis Type="Italic">Tamarix</Emphasis> (Tamaricaceae) hybrids: the dominant invasive genotype in southern Africa

Hybridization is regarded as a rapid mechanism for increasing genetic variation that can potentially enhance invasiveness. Tamarix hybrids appear to be the dominant genotypes in their invasions. Exotic Tamarix are declared invasive in South Africa and the exotic T. chinensis and T. ramosissima are known to hybridize between themselves, and with the indigenous T. usneoides. However, until now, it was not known which species or hybrid is the most prevalent in the invasion. With a biocontrol programme being considered as a way of suppressing the alien Tamarix populations, it is important to document the population genetic dynamics of all species in the region. This investigation sought to identify Tamarix species in southern Africa and their hybrids, describe their population structure, and reveal the geographic origin of the invasive species. To achieve this, nuclear Internal Transcribed Spacer (ITS) sequence data and the multilocus Amplified Fragment Length Polymorphisms (AFLPs) markers were used. Phylogenetic analysis and population genetic structure confirmed the presence of three species in South Africa (T. chinensis, T. ramosissima and T. usneoides) with their hybrids. The indigenous T. usneoides is clearly genetically distant from the alien species T. chinensis and T. ramosissima. Interestingly, the Tamarix infestation in South Africa is dominated (64.7 %) by hybrids between T. chinensis and T. ramosissima. The exotic species match their counterparts from their places of origin in Eurasia, as well as those forming part of the invasion in the US.     

Feasibility assessment for the classical biological control of <Emphasis Type="Italic">Tamarix</Emphasis> in Argentina

Saltcedars are woody plants in the genus Tamarix L. (Caryophyllales: Tamaricaceae) and are native to Eurasia and Africa. Several species have become invasive in the Americas, Australia and South Africa. In Argentina there are four species of Tamarix distributed in arid, semi-arid and coastal areas of most provinces. The taxonomic isolation of Tamarix spp. in Argentina, their widespread distribution, negative impact to natural areas and lack of impact from existing natural enemies all indicate that Tamarix is an ideal candidate for classical biological control in Argentina. Biological control of Tamarix spp. has been rapid and highly successful in the USA after the introduction of four Diorhabda spp. (Coleoptera: Chrysomelidae). Biological control of Tamarix spp. in Argentina could be implemented easily, rapidly, and at a low cost by utilizing the information developed in the USA.     

Ecophysiological responses of salt cedar (Tamarix spp. L.) to the northern tamarisk beetle (Diorhabdacarinulata Desbrochers) in a controlled environment

The northern tamarisk beetle (Diorhabda carinulata Desbrochers) was released in several western states as a biocontrol agent to suppress Tamarix spp. L. which has invaded riparian ecosystems; however, effects of beetle herbivory on Tamarix physiology are largely undocumented and may have ecosystem ramifications. Herbivory by this insect produces discoloration of leaves and premature leaf drop in these ecosystems, yet the cause of premature leaf drop and the effects of this leaf drop are still unknown. Insect herbivory may change leaf photosynthesis and respiration and may affect a plant’s ability to regulate water loss and increase water stress. Premature leaf drop may affect plant tissue chemistry and belowground carbon allocation. We conducted a greenhouse experiment to understand how Tamarix responds physiologically to adult beetle and larvae herbivory and to determine the proximate cause of premature leaf drop. We hypothesized that plants experiencing beetle herbivory would have greater leaf and root respiration rates, greater photosynthesis, increased water stress, inefficient leaf nitrogen retranslocation, lower root biomass and lower total non-structural carbohydrates in roots. Insect herbivory reduced photosynthesis rates, minimally affected respiration rates, but significantly increased water loss during daytime and nighttime hours and this produced increased water stress. The proximate cause for premature leaf drop appears to be desiccation. Plants exposed to herbivory were inefficient in their retranslocation of nitrogen before premature leaf drop. Root biomass showed a decreasing trend in plants subjected to herbivory. Stress induced by herbivory may render these trees less competitive in future growing seasons.     

The distribution and ecology of coastal species on roadsides

The invasion of roadsides by coastal species is a response to the use of de-icing salt. The most widespread species on Britain's roadsides, Puccinellia distans, occurs in the north and east of England and Wales. This distribution is due to the varying use and effects of de-icing salt which are related in turn to regional differences in climate and traffic density. On roadsides, seed dispersal is greater in the direction of traffic flow and the smaller seeds of Puccinellia distans are carried further than the larger seeds of Plantago maritima. Species on roadsides occur in distinct zones with the coastal species confined to the saline road margin. The causes of this zonation were investigated by examining seedling establishment in experimental plots. Seedlings were only able to establish in grass plots when the salinity was sufficiently high to kill the competing glycophytic grasses. On bare soil, the halophytes established and grew satisfactorily when salt was added but grew very poorly in the absence of salt. After two years Plantago maritima overcame this apparent requirement for salt on bare soil but Puccinellia distans and Spergularia marina did not.Acknowledgements: N. E. Scott was in receipt of a N.E.R.C. Studentship. The authors wish to thank the members of the B.S.B.I. and the Biological Records Office for information on species distribution.     

Can herbarium records be used to map alien species invasion and native species expansion over the past 100 years?

Aim To determine if the temporal and spatial pattern of alien plant invasion and native plant expansion can be observed using 100 years of herbarium data from Oklahoma, USA, and to eliminate herbarium collection biases in such analyses. Location Oklahoma, USA. Methods Using herbarium records from the Oklahoma Vascular Plants Database from 1903 to 2004, we reconstructed the spatial and temporal collection history of two alien invasive taxa (Lonicera japonica and Tamarix spp.) and three native expansive species (Ambrosia psilostachya, Amphiachyris dracunculoides and Juniperus virginiana). To compare the overall collecting trend, groups of native non‐expansive taxa were selected as counterparts. We recorded the year of the first collection in each township in Oklahoma for all taxa. The cumulative number of occupied townships was log‐transformed, plotted against time and modelled with linear regression. The slope of the linear regression represented collection trend over time for the non‐expansive counterpart group. However, for the invasive and expansive species, the regression slope represented the collection effort plus the invasion or expansion rate. We calculated the proportion of invasive and expansive species to non‐expansive species by dividing the cumulative number of townships for each invasive or expansive species by the cumulative number of townships occupied by the counterpart group (proportion curve). Results Maps of the collection records of invasive and expansive taxa illustrated no discernible spatial invasion or expansion pattern. The slopes of the linear regression for alien invasive taxa were significantly steeper than those of their associated native non‐expansive counterparts, indicating an increase in abundance. Juniperus virginiana, L. japonica and Tamarix spp. exhibited one or more periods during which they were collected at a disproportionately higher rate than their native non‐expansive counterparts. Main conclusions Patterns of species invasion and expansion in Oklahoma were detected using techniques developed for regions with longer collecting plant histories. The proportion curve analysis eliminated some biases inherent in herbarium data by reducing the effect of collecting effort. Both the regression model and proportion curve analyses illustrate the temporal invasion patterns of alien invasive species. The native species did not show a clear expansion pattern. The information found in recently established herbaria may not be sensitive enough to detect the increase in abundance of native species.