The alien flora of Europe: a taxonomic and biogeographic review

Abstract. A geographic and taxonomic overview of the non-indigenous plant species of Europe, based on the ‘Flora Europaea’ is given. The flora of Europe includes 1568 species which have either expanded their ranges within Europe under human influence (naturalized European species) or are of non-European origin (exotic species). The latter group consists of 580 species (37%) which form a diverse group in terms of their taxonomic composition and geographic origin. The exotics are represented by 113 families, of which the Compositae, Rosaceae and Gramineae are most important. The ratio of species to families is low. Most exotic species in Europe originate from the Americas and Asia. Countries of southern Europe have a higher relative number of exotics in their non-native flora than northern ones. The species-range size distribution differs between naturalized European and exotic species; the latter are on average more widespread than the naturalized.     

Pachybrachis (Coleoptera,Chrysomelidae, Cryptocephalinae) of Eastern Canada

Seventeen Pachybrachis species occurring in eastern Canada [Ontario (ON), Québec (QC), New Brunswick (NB), Nova Scotia (NS), and Prince Edward Island (PE)] are treated by the authors. Two new national records were discovered, both from southernmost Ontario: P. cephalicus Fall and P. luctuosus Suffrian. Four species were new provincial records: P. cephalicus (ON), P. luctuosus (ON, QC), P. obsoletus Suffrian (NB), P. peccans (PE). A fully illustrated key to the Pachybrachis of eastern Canada is provided and supported with extensive photographs, distribution maps, and plant associations. Three species were distributed from southern Ontario into at least one province in the Maritimes (P. nigricornis (Say), P. obsoletus Suffrianand P. peccans Suffrian). Six species were distributed along the shores of the Great Lakes (Erie, Michigan, and Ontario) and rivers (Ottawa, Saguenay and St. Lawrence), but unknown from central and northern ON and QC (P. bivittatus (Say), P. hepaticus hepaticus (F. E. Melsheimer), P. othonus othonus (Say), P. pectoralis (F. E. Melsheimer), P. spumarius Suffrianand P. trinotatus (F. E. Melsheimer)). Seven species were rare, five being found exclusively from southern ON (P. calcaratus Fall, P. cephalicus, P. luridus (Fabricius), P. subfasciatus (J. E. LeConte)and P. tridens (F. E. Melsheimer)), with two having, in addition, a disjunct population in QC (P. atomarius (F. E. Melsheimer)and P. luctuosus). One species was found to be the northern most extension of an eastern United States (US) distribution into the eastern townships of QC (P. m-nigrum (F. E. Melsheimer)). There were no Pachybrachis that could be considered arctic, subarctic, or boreal species; no specimens were found from Labrador and Newfoundland, and all species had southern affinities. Pachybrachis atomarius, P. calcaratus, P. luridus, P. subfaciatus, and P. tridens, not seen over the last 30–70 years, may be extirpated from eastern Canada.     

Exotic plant species in a C4-dominated grassland: invasibility, disturbance, and community structure

We used data from a 15-year experiment in a C₄-dominated grassland to address the effects of community structure (i.e., plant species richness, dominance) and disturbance on invasibility, as measured by abundance and richness of exotic species. Our specific objectives were to assess the temporal and spatial patterns of exotic plant species in a native grassland in Kansas (USA) and to determine the factors that control exotic species abundance and richness (i.e., invasibility). Exotic species (90% C₃ plants) comprised approximately 10% of the flora, and their turnover was relatively high (30%) over the 15-year period. We found that disturbances significantly affected the abundance and richness of exotic species. In particular, long-term annually burned watersheds had lower cover of exotic species than unburned watersheds, and fire reduced exotic species richness by 80–90%. Exotic and native species richness were positively correlated across sites subjected to different fire (r = 0.72) and grazing (r = 0.67) treatments, and the number of exotic species was lowest on sites with the highest productivity of C₄ grasses (i.e., high dominance). These results provide strong evidence for the role of community structure, as affected by disturbance, in determining invasibility of this grassland. Moreover, a significant positive relationship between exotic and native species richness was observed within a disturbance regime (annually burned sites, r = 0.51; unburned sites, r = 0.59). Thus, invasibility of this C₄-dominated grassland can also be directly related to community structure independent of disturbance. Received: 9 February 1999 / Accepted: 12 May 1999     

Reintroduction of Nassella pulchra to California coastal grasslands: Effects of topsoil removal,plant neighbour removal and grazing

Question: What is the most appropriate combination of treatments to reintroduce Nassella pulchra, a perennial bunchgrass, into degraded mediterranean coastal grasslands? Location: Central coast of California, USA. Methods: N. pulchra was sown from seeds and transplanted into a degraded grassland in a multi‐factorial experiment testing the effects of (1) two grazing intensities (lightly grazed by native mammal species or ungrazed); (2) topsoil removal and (3) reduction of plant neighbours. The experiment was carried out on two types of surrounding vegetation (exotic annual grasses and exotic forbs). Results: Topsoil removal greatly enhanced establishment from seeds and transplant survival, mainly because it reduced the exotic vegetation and thus reduced competition. While removing neighbours was essential when topsoil was left intact, it had a negative effect on N. pulchra when surrounding species included exotic forbs (Brassica spec, and Asteraceae) at low density (after topsoil removal). Moderate grazing by native mammals (deer, rabbits and gophers) did not affect N. pulchra. Conclusion: Our results suggest that seeding after topsoil has been removed is a promising method to reintroduce N. pulchra to highly degraded sites where there is little to no native seed bank.     

Deposition of exotic bird-dispersed seeds into three habitats of a fragmented landscape in the northeastern United States

Dispersal by frugivorous birds facilitates invasion by many exotic plants. We measured the seed rain of ornithochorous native and exotic plants at three habitats of a fragmented landscape of the northeastern United States for 1 year. We studied maple-beech forests, old fields, and abandoned conifer plantations. Across all sites we collected 2,196 ornithochorous seeds, including seeds from six exotic species and 10 native species. The majority (90%) of collected seeds were from exotic species. Seed dispersal was broadly similar among habitats, though seed rain of exotic species was higher in old fields than forested habitats. Seed rain was not strongly influenced by artificial perches for most species. However, seeds of exotic species were more commonly found in traps under an artificial perch in old fields. Seed rains for the exotic Elaeagnus umbellata, Rhamnus cathartica, and Rosa multiflora were positively associated with local density of mature plants. Seed rain of R. cathartica was positively associated with abundance of seedlings but not saplings, suggesting that post-dispersal mortality was important. Seed dispersal of the exotic Lonicera spp. was high in all habitats, accounting for 66% of all seeds collected. With the exception of Lonicera spp., seed rain of common exotic invaders was affected by the abundance of seed sources, and these species might be effectively controlled by elimination of local fruiting plants. Fruits of Lonicera morrowii, which has extensively invaded our area, are apparently a common component in the diet of frugivores.     

California native and exotic perennial grasses differ in their response to soil nitrogen,exotic annual grass density,and order of emergence

Early emergence of plant seedlings can offer strong competitive advantages over later-germinating neighbors through the preemption of limiting resources. This phenomenon may have contributed to the persistent dominance of European annual grasses over native perennial grasses in California grasslands, since the former species typically germinate earlier in the growing season than the latter and grow rapidly after establishing. Recently, European perennial grasses have been spreading into both non-native annual and native perennial coastal grass stands in California. These exotic perennials appear to be less affected by the priority effects arising from earlier germination by European annual grasses. In addition, these species interactions in California grasslands may be mediated by increasing anthropogenic or natural soil nitrogen inputs. We conducted a greenhouse experiment to test the effects of order of emergence and annual grass seedling density on native and exotic perennial grass seedling performance across different levels of nitrogen availability. We manipulated the order of emergence and density of an exotic annual grass (Bromus diandrus) grown with either Nassella pulchra (native perennial grass), Festuca rubra (native perennial grass), or Holcus lanatus (exotic perennial grass), with and without added nitrogen. Earlier B. diandrus emergence and higher B. diandrus density resulted in greater reduction in the aboveground productivity of the perennial grasses. However, B. diandrus suppressed both native perennials to a greater extent than it did H. lanatus. Nitrogen addition had no effect on the productivity of native perennials, but greatly increased the growth of the exotic perennial H. lanatus, grown with B. diandrus. These results suggest that the order of emergence of exotic annual versus native perennial grass seedlings could play an important role in the continued dominance of exotic annual grasses in California. The expansion of the exotic perennial grass H. lanatus in coastal California may be linked to its higher tolerance of earlier-emerging annual grasses and its ability to access soil resources amidst high densities of annual grasses.     

The effect of soil nitrogen on competition between native and exotic perennial grasses from northern coastal California

The invasion of European perennial grasses represents a new threat to the native coastal prairie of northern California. Many coastal prairie sites also experience anthropogenic nitrogen (N) deposition or increased N availability as a result of invasion by N-fixing shrubs. We tested the hypothesis that greater seedling competitive ability and greater responsiveness to high N availability of exotic perennial grasses facilitates their invasion in coastal prairie. We evaluated pairwise competitive responses and effects, and the occurrence of asymmetrical competition, among three common native perennial grasses (Agrostis oregonensis, Festuca rubra, and Nassella pulchra) and three exotic perennial grasses (Holcus lanatus, Phalaris aquatica, and Festuca arundinacea), at two levels of soil N. We also compared the root and shoot biomass and response to fertilization of singly-grown plants, so we could evaluate how performance in competition related to innate plant traits. Competitive effects and responses were negatively correlated and in general varied continuously across native and exotic species. Two exceptions were the exotic species Holcus, which had large effects on neighbors and small responses to them, and competed asymmetrically with all other species in the experiment, and the native grass Nassella, which had strong responses to but little effect on neighbors, and was out-competed by all but one other species in the experiment. High allocation to roots and high early relative growth rate appear to explain Holcus’s competitive dominance, but its shoot biomass when grown alone was not significantly greater than those of the species it out-competed. Competitive dynamics were unaffected by fertilization. Therefore, we conclude that seedling competitive ability alone does not explain the increasing dominance of exotic perennial grasses in California coastal prairie. Furthermore, since native and exotic species responded individualistically, grouping species as ‘natives’ and ‘exotics’ obscured underlying variation within the two categories. Finally, elevated soil N does not appear to influence competition among the native and exotic perennial grasses studied, so reducing soil N pools may not be a critical step for the restoration of California coastal prairie.     

Afforestation causes changes in post‐fire regeneration in native shrubland communities of northwestern Patagonia,Argentina

Question: What are the effects of fire in native shrubland communities and in pine plantations established in these shrublands? Location: Northern Patagonia, Argentina. Methods: We surveyed four sites in Chall‐Huaco valley, located in northwest Patagonia. Each site was a vegetation mosaic composed of an unburned Pinus ponderosa plantation, a plantation burned in 1996, and an unburned matorral and a matorral burned by the same fire. We recorded the cover of all vascular plant species. We also analysed species richness, total cover, proportion of exotic species, abundance of woody species and herb species, cover of exotic species, abundance of woody and herb species and differences in composition of species. For both shrubs and tree species we recorded the main strategy of regeneration (by resprouting or by seed). Results: We found that fire had different effects on native matorral and pine plantations. Five years after fire, plantations came to be dominated by herbs and exotic species, showing differences in floristic composition. In contrast, matorral communities remained very similar to unburned matorral in terms of species richness, proportion of woody species, and herb species and proportion of exotics. Also, pine plantations were primarily colonized by seedlings, while matorrals were primarily colonized by resprouting. Conclusions: Matorrals are highly fire resilient communities, and the practice of establishing plantations on matorrals produces a strong reduction in the capacity of matorral to return to its original state. The elimination of shrubs owing to the effect of plantations can hinder regeneration of native ecosystems. Burned plantations may slowly develop into ecosystems similar to the native ones, or they may produce a new ecosystem dominated by exotic herbs. This study shows that plantations of exotic conifers affect native vegetation even after they have been removed, as in this case by fire.     

Using integral projection models to compare population dynamics of four closely related species

Demographic processes, such as survival, growth, and reproduction, can inform us about invasion risk, extinction risk, and trade-offs in life history strategies. The population dynamics of four Amaranthaceae species in southern Illinois, USA were examined using integral projection models (IPMs) to determine whether vital rates reflect life history among these closely related species. Two of the species, Amaranthus palmeri and Amaranthus tuberculatus, are summer annuals and considered to be some of the most problematic agricultural weeds in the US Midwest. Achyranthes japonica is a relatively new invasive exotic species that primarily inhabits forests. Iresine rhizomatosa, is an endangered species in the study area, which also inhabits forests. Two populations of each species were studied from 2012 to 2014 in which height of individuals were measured and used as the state variable in the IPMs. The Amaranthus species and Achyranthes japonica had an estimated population growth rate >1, projecting increases in population size. By contrast, λ was <1 for I. rhizomatosa, projecting a decline in population size demonstrating its endangered status. Germination rates and seed viability were dependent on species and varied over time. Elasticity analyses showed that survival and growth contributed most to λ for the perennial species; whereas, for the annual species population dynamics were driven primarily by fecundity. Overall, Achyranthes japonica and the Amaranthus species show similar trends in demographic processes that align with their invasive nature and not with their life histories. Furthermore, this study demonstrates that more research on the competitive nature of Achyranthes japonica is needed.     

Fruit Mineral Contents of Six Wild Species of the North Andean Patagonia, Argentina

The fruit mineral contents (K, Ca, Ba, Br, Zn, Co, Cr, Fe, Na, Rb, Cs, and Sr) of four native and two exotic naturalized shrubs growing in different areas of the Andean Patagonian region of Argentina were investigated. Native species Berberis darwinii, Berberis microphylla (Berberidaceae), Aristotelia chilensis (Elaeocarpaceae) and Ribes magellanicum (Saxifragaceae) produce small berries while the fruits of the exotic species Rosa rubiginosa and Rosa canina (Rosaceae) are aggregates of aquenes. They are used to prepare jams, tea, liquors, and ice creams. Native shrub fruits had higher content of Br, Zn, Co, Cr, Fe, Mo, and Na than those of the exotic naturalized species. Rosa species showed the highest contents of Ca and Ba in their fruits (the mean content doubled those of the native plant fruits). The fruit nutrient content found in the studied species was similar or higher than other values reported for fruits of temperate and tropical species in the world.     

Land use history, hurricane disturbance, and the fate of introduced species in a subtropical wet forest in Puerto Rico

Tropical forests are suffering from increasing intensities and frequency of disturbances. As a result, non-native species accidentally introduced or intentionally planted for farming, plantations, and ornamental purposes may spread and potentially invade undisturbed native forest. It is not known if these introduced species will become invasive, as a result of recurrent natural disturbances such as hurricanes. Using data from three censuses (spanning 15 years) of a 16-ha subtropical wet forest plot, we investigated the impact of two hurricanes on populations of plant species that were planted in farms and plantations that were then abandoned and from the natural spread of species introduced into Puerto Rico in the past. The populations of four species (Citrus paradis, Mangifera indica, Musa sp., and Simarouba glauca) changed little over time. Six species (Artocarpus altilis, Calophyllum calaba, Genipa americana, Hibiscus pernambucensis, Syzygium jambos, and Swietenia macrophylla) declined between the first two censuses after Hurricane Hugo, then increased again in Census 3 after Hurricane Georges. Spathodea campanulata gradually increased from census to census, while Coffea arabica declined. These introduced species represent only a small part of the forest basal area and few show signs of increasing over time. The number of stems per plant, new recruits, and the growth rates of these introduced species were within the ranges of those for native plant species. The mortality rates over both census intervals were significantly lower for introduced species (<5% year⁻¹) than for native ones (15% year⁻¹). Many new recruits established after Hurricane Hugo (prior to this study) had opened the forest canopy and these high mortality rates reflect their death as the canopy recovered. Only Swietenia macrophylla and Syzygium jambos showed an increase in stem numbers in the closed canopy area of forest that had suffered limited human disturbance in the past. A future increase in frequency of disturbance may enable greater stem numbers of introduced species to establish, while lower-mortality rates compared to native species, may allow them to persist during inter-hurricane intervals. An increase in the population of introduced species, especially for those that grow into large trees, may have an impact on this tropical forest in the future.     

Dominance of non-native riparian trees in western USA

Concern about spread of non-native riparian trees in the western USA has led to Congressional proposals to accelerate control efforts. Debate over these proposals is frustrated by limited knowledge of non-native species distribution and abundance. We measured abundance of 44 riparian woody plants at 475 randomly selected stream gaging stations in 17 western states. Our sample indicates that Tamarix ramosissima and Elaeagnus angustifolia are already the third and fourth most frequently occurring woody riparian plants in the region. Although many species of Tamarix have been reported in the region, T. ramosissima (here including T. chinensis and hybrids) is by far the most abundant. The frequency of occurrence of T. ramosissima has a strong positive relation with the mean annual minimum temperature, which is consistent with hypothesized frost sensitivity. In contrast the frequency of occurrence of E. angustifolia decreases with increasing minimum temperatures. Based on mean normalized cover, T. ramosissima and E. angustifolia are the second and fifth most dominant woody riparian species in the western USA. The dominance of T. ramosissima has been suspected for decades; the regional ascendance of E. angustifolia, however, has not previously been reported.     

Exotic ladybirds for biological control of herbivorous insects – a review

Since the late 19th century, exotic ladybirds (Coleoptera: Coccinellidae) have been used extensively for suppressing herbivorous insects of economic importance. In recent decades, the introduction of non‐native biological control (BC) agents has been greatly limited due to the awareness of the potential non‐target effects of introductions. Nonetheless, recent episodes of biological invasions of economically important pests have raised the need to carefully consider whether the expected benefits of pest control go beyond the possible environmental risks of introduction. To better understand the factors that contributed to successful BC programs, here we review the literature behind classical and augmentative BC using exotic ladybirds. Additionally, by means of case studies, we discuss the BC efficacy of selected exotic species, e.g., Coccinella septempunctata L., Harmonia axyridis (Pallas), and Hippodamia variegata (Goeze), and their position within the communities of predators in the introduced areas of USA, Canada, and Chile. In Europe, much of the research on exotic ladybirds has been conducted on the undesired impact of H. axyridis. Therefore, we summarize the risk assessment data for this species and review the field research investigating the ecological impact on European aphidophagous predators. According to the BIOCAT database of classical BC programs, 212 ladybird species belonging to 68 genera have been released in about 130 years of BC activity, with 14.6% of introductions having resulted in partial, substantial, or complete control of the target pest. However, because post‐release evaluation of establishment and BC success has not always been conducted, this rate could underestimate the successful cases. Among other factors, ladybird establishment and pest suppression mostly depend on (1) intrinsic factors, i.e., high voracity, synchronized predator‐prey life cycle, and high dispersal ability, and (2) extrinsic factors, i.e., adaptability to the new environment and landscape composition. This review contributes to improved understanding of ladybirds as exotic BC agents.     

Detection of European Peristenus digoneutis loan in mirid populations in southern Ontario

Lygus spp. Hahn (Hemiptera: Miridae) are serious pests of a wide variety of economically important crops in North America. A European parasitoid, Peristenus digoneutis Loan (Hymenoptera: Braconidae), released in the northeastern USA for the biological control of Lygus, has successfully established in nine states and in eastern Canada, including southeastern Ontario, southern Quebec and Nova Scotia. To determine the extent to which P. digoneutis has dispersed and established in mirid populations in Ontario, a single-step multiplex PCR assay, designed to differentiate European species P. digoneutis and P. relictus from native North American Peristenus species, was used to identify parasitoid larvae dissected from field-collected Miridae. A total of 222 parasitoid larvae were analysed with the multiplex assay. Most (172) were identified as members of the P. pallipes Curtis complex; however, three from the Niagara region were identified as P. digoneutis. Specimens that did not amplify using the Peristenus multiplex PCR assay were screened with a PCR primer set designed to detect Leiophron (Hymenoptera: Braconidae) species, and most were confirmed to be Leiophron larvae. This study confirms the presence of P. digoneutis species in southern Ontario, and is another example of the utility of molecular methods for the detection of newly introduced or dispersed parasitoids. The presence of exotic P. digoneutis in southern Ontario may expedite future releases to augment already-established populations. Following such releases, the Peristenus multiplex PCR assay will be a useful component of post-release studies to evaluate the success of the biological control programme for Lygus plant bugs in Canada.     

Arthropod diversity of exotic vs. native Robinia species in northern Arizona

1 Arthropods were collected on native locust, Robinia neomexicana A. Gray, and exotic Robinia pseudoacacia L. in northern Arizona over a 2‐year period to determine the number of arthropod species and number of individuals present. 2 More arthropod species were found on the native (251) than on the exotic Robinia (174). 3 Greater species diversity was likewise found on the native than the exotic. The five most numerous insects collected each year accounted for 81% to 91% of the total number collected on the exotic and native Robinia in 1997 and 1998. Only 12 species occurred on both the native and exotic Robinia in both years. 4 These findings are discussed in the context of using exotic trees in plantations and ecological theory regarding rates of arthropod species accumulation on exotic hosts.     

The fate of an intentional introduction of Formica lugubris to North America from Europe

Red wood ants (Formica s.str.) are not prevalent in the forests of North America, but commonly occur in conifer and mixed conifer forests in northern Europe and Asia. In 1971, a European red wood ant species, Formica lugubris, was intentionally established in a 35‐year‐old predominantly mixed conifer plantation approximately 30 km north of QC, Canada. The purpose of its introduction was to evaluate the potential of this species as a biological control agent against conifer‐defoliating Lepidoptera species. This red wood ant introduction was monitored periodically for about 5 years after establishment, but its long‐term fate has not been reported. We visited this field site in 2005 and found that this species was well established, and we could locate some of the nests that resulted from the original release. We mapped and measured over 100 nests around the site of original release, which ranged from 5 cm in height to over 1 m. We estimated the population of introduced ants to have grown to over 8 million in the last 34 years. Significant clustering of nests suggests that these nests may be one supercolony. F. lugubris has become a dominant understory arthropod in this mixed forest, and is likely to have ecological impacts, including effects at the community and ecosystem level.     

Inter-regional hybrids of native and non-native Centaurea sulphurea inherit increased competitive ability from the non-natives

Background: Exotic species can rapidly develop adaptations to their non-native regions, such as increased size and competitive ability. Although these traits are believed to be responsible for invasive success, some non-invasive exotic species display them too. This suggests that increased size and competitive ability might be necessary but not sufficient to turn an exotic into a successful invader.

Aim: To assess size and competitive ability of the exotic non-invasive weed Centaurea sulphurea, and of inter-regional hybrids between native and non-native populations.

Methods: We experimentally produced a cohort of C. sulphurea individuals from the native range of the species in Spain, from its non-native range in California, as well as hybrids between the two regions. We grew these plants in pots in competition with the grass Bromus hordeaceus, or alone in control pots.

Results: Individuals from California were larger and better competitors than individuals from Spain. Furthermore, inter-regional hybrids showed competitive responses similar to that of individuals from California.

Conclusions: Our results confirm that increased competitive ability might be more frequent than previously thought among introduced species, since it can be detected in at least some exotic non-invasive species. They also illustrate the importance of maternal effects, how locally adapted traits are conserved and spread in the non-native ranges of exotic species, and suggest that plant size and competitive ability are not directly associated in this species.     

Northern pike (<Emphasis Type="Italic">Esox lucius</Emphasis>) in a warming lake: changes in population size and individual condition in relation to prey abundance

The spring-fed Comal River in Texas, USA, has been impounded and channelized resulting mainly in a lentic environment with four headwater spring runs. We sampled two spring runs (lotic sites) and two lentic sites seasonally from April 2001 through April 2002 to assess (1) co-occurrence of native and exotic snail species (as determined by interspecific association), (2) the importance of habitat conditions in structuring relationships among these species, and (3) the distribution of snails infected with exotic trematode parasites. Three exotic and four endemic species of aquatic snails were collected, but only Elimia comalensis (Prosobranchia: Pleuroceridae, native), Melanoides tuberculatus, and Tarebia granifera (Prosobranchia: Thiaridae, exotic) were in sufficient densities for further analyses. Tarebia granifera was positively associated with both M. tuberculatus (χ² = 18.5, P < 0.001) and E. comalensis (χ² = 7.3, P < 0.01), although the co-occurrence between the two exotics was much stronger. Melanoides tuberculatus and E. comalensis exhibited a strong, negative association (χ² = 10.9, P < 0.001). The weaker co-occurrence between E. comalensis with the thiarids appeared to be driven by differences in habitat use by the thiarids and native E. comalensis. In lentic habitats, densities of M. tuberculatus and T. granifera were similar but differed significantly from E. comalensis whose densities were 200 times less than the exotic snails. In lotic spring runs, densities of T. granifera and E. comalensis were similar, but differed significantly from M. tuberculatus whose densities were 10 times fewer. Lower densities of M. tuberculatus and T. granifera in habitat conditions common to the spring runs may explain why exotic snail interactions were less with the native E. comalensis than with each other. The native snail, E. comalensis, was not infected with any trematodes, while 6.1% of M. tuberculatus and 4.8% of T. granifera were infected with exotic trematodes. Distributions of infected snails were aggregated; such that most infected snails were found in lentic habitats with silt substrates and moderate to high levels of detritus. Continued declines in spring-flows due to aquifer withdrawals and droughts will increase lentic habitats that may lead to increased densities of T. granifera and M. tuberculatus and their concomitant parasites. Handling editor: K. Martens     

Exotic plant invasion in the highly degraded upland landscape of Hong Kong, China

Plant communities in the continental tropics have suffered less from exotic plant invasions than their oceanic island counterparts. Most studies have focused on near-pristine communities. By contrast, we examine the resistance of semi-natural continental plant communities in Hong Kong, which have been suffering from chronic and massive human impacts. We compiled a list of all naturalized non-native species recorded in Hong Kong and then sampled the plant communities for exotic species along roadsides, a stream through semi-natural vegetation, and in semi-natural vegetation away from both roads and streams on Tai Mo Shan, Hong Kong’s highest peak (957 m). Similar surveys were repeated in other areas of Hong Kong. More than 162 naturalized exotic plant species have been recorded in Hong Kong. On Tai Mo Shan, 29 exotic species were recorded in roadside vegetation, with the diversity but not percentage cover declining significantly with altitude. Fifteen exotic species were found along the stream, including two not found along the roadside. Only six exotic species were found away from roads and streams, all in unshaded areas disturbed by feral cattle. In all surveys, no exotics were found in closed woody vegetation or in open areas without feral cattle, except for one species. The shade-tolerant tropical Asian tree Syzygium jambos was found invading along some streamsides without anthropogenic disturbance. Despite centuries of massive human impacts, exotic plant invasions in Hong Kong are still largely confined to habitats that suffer from chronic human disturbance. Feral cattle promote invasion where people are absent, but this problem still seems potentially reversible. Only Syzygium jambos is of possible current conservation concern.