Mutualisms with the wreckage of an avifauna: the status of bird pollination and fruit dispersal in New Zealand

Worldwide declines in bird numbers have recently renewed interest in how well bird?plant mutualisms are functioning. In New Zealand, it has been argued that bird pollination was relatively unimportant and bird pollination failure was unlikely to threaten any New Zealand plants, whereas dispersal mutualisms were widespread and in some cases potentially at risk because of reliance on a single large frugivore, the kereru (Hemiphaga novaeseelandiae). Work since 1989, however, has changed that assessment. Smaller individual fruits of most plant species can be dispersed by mid-sized birds such as tui (Prosthemadera novaezelandiae) because both fruits and birds vary in size within a species. Only one species (Beilschmiedia tarairi) has no individual fruits small enough for this to occur. Germination of 19 fleshy-fruited species, including most species with fruits >8 mm diameter, does not depend on birds removing the fruit pulp. The few studies of fruit removal rates mostly (7 out of 10) show good dispersal quantity. So dispersal is less at risk than once thought. In contrast, there is now evidence for widespread pollen limitation in species with ornithophilous flowers. Tests on 10 of the 29 known native ornithophilous-flowered species found that in 8 cases seed production was reduced by at least one-third, and the pollen limitation indices overall were significantly higher than the global average. Birds also frequently visit flowers of many other smaller-flowered native species, and excluding birds significantly reduced seed set in the three species tested. So pollination is more at risk than once thought. Finally, analyses of both species numbers and total woody basal area show that dependence on bird pollination is unexpectedly high. Birds have been recorded visiting the flowers of 85 native species, representing 5% of the total seed-plant flora (compared with 12% of those with fleshy fruit) and 30% of the tree flora (compared with 59% with fleshy fruit). A higher percentage of New Zealand forest basal area has bird-visited flowers (37% of basal area nationally) than fleshy fruit (31%). Thus, bird pollination is more important in New Zealand than was realised, partly because birds visit many flowers that do not have classic ?ornithophilous? flower morphology.     

Breeding System and pollination of selected plants endemic to Juan Fernandez Islands

We conducted field studies on the Juan Fernández Islands flora on the breeding system of 25 endemic species from 17 families. We recorded data on flower features, pollen and ovule number, pollen/ovule ratio, pollen size, self-compatibility, floral visitors, and pollination. Flowers are mostly hermaphrodite, inconspicuous, small, and green. Six species are dioecious. Over 80% of the cosexual species are self compatible. However, many species are dichogamous (mostly protandrous); thus, even the self-compatible species may require pollen transfer. Selfing through geitonogamy seems to be the most common system, and several species express mixed breeding systems. Floral visitors are uncommon to rare, except for two hummingbird species (one native and one endemic) that visit five species we studied. In more than 300 h of observation of flowers over three field seasons, we detected only 23 native insect visits representing ten species (Diptera, Lepidoptera, and Coleoptera). One species each of an introduced ant and an introduced bee were also observed on some flowers, all near the single human settlement of San Juan Bautista. Wind directly moving pollen, or indirectly via shaking the flowers, is the most important pollen distribution mechanism. The majority of the wind-pollinated species bear some typical anemophilous features, but also others not characteristic of wind pollination, that presumably represent the condition of their biotically pollinated ancestors. Floral features often reflect ancestral reproductive systems, so floral biology studies of oceanic islands in particular must be done with cognizance of presumed ancestral forms, because the observed characters can be misleading.     

Sex ratio and gender stability in the dioecious plants of Israel

The hypothesis that in dioecious plants genders are usually stable and that the sex ratio is 1:l was tested in the dioecious species of Israel. The flora of Israel comprises some 2500 wild species, among which 43 are dioecious, one gynodioecious and one androdioecious. Forty-one out of these 45 species were examined during 1995-97. These species were examined in 120 populations comprising 7019 individuals, at 81 different localities. The majority of the species (33/41) manifested the expected 1:1 sex ratio. Of the eight species deviating from this ratio, a significant bias was repeatedly scored in five of them. Three other species of the willow family manifested sometimes strongly biased sex ratios or even monomorphic (unisexual) stands, apparently as a result of extensive vegetative cloning. The stability of the male and female morphs was also examined in the 41 species. Only eight individuals (from seven species) out of the 7019 individuals surveyed (0.1%) were found to be bisexual. In addition, in 27 populations belonging to 17 species, 909 males were tagged and revisited in fruiting time, and only three of them (0.3%) were found to set fruits. In addition not a single case of gender reversal between seasons was found. The field survey suggests that in dioecious plants sex determination and expression are usually effectively maintained. The very slight deviations from the 1:1 sex ratio and from the distinct male and female morphs suggest that sexual dimorphism in dioecious plants is usually stabile.     

Threatened pollination systems in native flora of the Ogasawara (Bonin) Islands

• Background and Aims Various alien species have been introducedto the Ogasawara Islands (Japan). A survey was made investigatingwhether the native pollination systems fit an ‘islandsyndrome’ (biasing the flora to dioecy, with subdued,inconspicuous flowers) and whether alien species have disruptedthe native pollination network. • Methods Flower visitors and floral traits were determinedin the field (12 islands) and from the literature. Associationsamong floral traits such as sexual expression, flower colourand flower shape were tested. • Key Results Among the 269 native flowering plants, 74·7% are hermaphroditic, 13·0 % are dioecious and 7·1% are monoecious. Classification by flower colour revealed that36·0 % were white, 21·6 % green and 13·8% yellow. Woody species (trees and shrubs) comprised 36·5% of the flora and were associated with dioecy and white flowers.Solitary, endemic small bees were the dominant flower visitorsand visited 66·7 % of the observed species on satelliteislands where the native pollination networks are preserved.In contrast to the situation on the satellite islands, introducedhoneybees were the most dominant pollinator (visiting 60·1% of observed species) on the two main islands, Chichi-jimaand Haha-jima, and had spread to satellite islands near Chichi-jimaIsland. • Conclusions The island syndrome for pollination systemsin the Ogasawara Islands was evident in a high percentage ofdioecious species, the subdued colour of the native flora andsolitary flower visitors on satellite islands. The shape andcolour adaptations of several flowers suggested native pollinationniches for long-proboscis moths and carpenter bees. However,the domination and expansion of introduced honeybees have thepotential for disruption of the native pollination network inthe two main, and several satellite, islands of the OgasawaraIslands.     

Alien and native species in Central European urban floras: a quantitative comparison

The paper provides quantitative information on the occurrence of alien species in Central European cities and analyses factors determining the richness of alien and native floras in this habitat type. Data for 54 cities (25 Polish, 24 German, 4 Czech and 1 Austrian) were gathered, and the representation of archaeophytes (i.e. aliens introduced before 1500 ad ), neophytes (introduced after that date) and native species was expressed. In an average city there were 87.4 archaeophytes (15.2% of the city flora) and 172.4 neophytes (25.2%) giving a total of 259.7 for alien species (40.3%). The number of native species averaged 386.5. The numbers of species in each category of immigration status increased significantly with city size. For neophytes, the species-area relationship showed a higher slope (0.49) on log–log axes than for both archaeophytes (0.16) and native species (0.30). Not only the number, but also the relative contribution of neophytes to the total flora, increased with city size, indicating that neophytes are the group which are most closely associated with human activity. On the other hand, archaeophytes were better represented in smaller cities, as they were confined to rural environment. A step-wise multiple regression was used to test for environmental variables acting as significant predictors, and explained between 40 and 65% of variation in the species numbers for particular categories of immigration status, providing the best fit for neophytes. City size was the best predictor for each characteristic, except of the proportion of total aliens, where the percentage of explained variability was low (8.2%), with latitude being the only significant predictor. Temperature was another highly significant predictor for the number of archaeophytes and total aliens, reflecting the origin of aliens in warmer areas. There was an effect of region on some flora characteristics. Polish cities had significantly higher proportion of archaeophytes and of total aliens than German cities. It is concluded that the occurrence of native and alien species in urban floras follows rather different pattern.     

DIOECISM AND ITS CORRELATES IN THE CAPE FLORA OF SOUTH AFRICA

Floral sexuality is characterized for the flora of the Cape region of South Africa. Among angiosperms (N = 8,497), monostylous hermaphroditism constitutes the largest proportion of species with 77.7%, followed by gynomonoecism (7.1%), dioecism (6.6%), andromonoecism (4.0%), monoecism (2.6%), heterostylous hermaphroditism (1.9%), polygamomonoecism (0.01%) and polygamodioecism (0.01%). The incidence of dioecism is significantly higher (P < 0.001) at the species level for two smaller floras within the Cape flora that consist mostly of species-rich fynbos vegetation (Cape of Good Hope Nature Reserve: 11.6% [N = 1,349], Cape Hangklip: 9.7% [N = 1,046]). The relatively high incidence of dioecism in all three floras compared to other temperate floras is due to the large number of Restionaceae species. Excluding these rush-like plants, dioecism at the species level is only 3.0% for the Cape flora, 4.2% for the Cape Hangklip flora and 3.6% for the Cape of Good Hope flora. At the generic level there are no significant differences in the incidence of dioecism between the three floras. Among dioecious species and genera in the Cape flora, there is a higher than expected incidence of wind pollination, fleshy fruits and nonwoodiness when each variable is analyzed independently. The association between nonwoodiness and dioecy has not been reported for other floras and is due to the large proportion of nonwoody wind-pollinated Restionaceae. If this family is excluded from the analysis, dioecy becomes associated with biotic pollination, fleshy fruits, and woodiness. Interactions among the variables themselves as well as with dioecism indicate the need for a joint analysis of the variables. At the genus level, analyses reveal the following: 1) There is a higher than expected incidence of wind pollination among dioecious plants with dry fruits. If the Restionaceae are excluded from the analysis, wind pollination is more common than expected only for woody plants with dry fruits. 2) Dioecious plants that are biotically pollinated have a significantly higher incidence of fleshy fruits than expected. 3) Dioecious plants with dry fruits that are biotically pollinated have a higher than expected incidence of woodiness. If the Restionaceae are excluded from the analysis, woodiness is more common than expected among both nonwind and wind-pollinated dioecious genera.     

The Physiology of Sexual Reproduction in Lunularia cruciata (L.) Dum.

Abstract

Since 1794 about 1.3 species per year have been added to the known bryophyte flora of Oxfordshire.This is chiefly due to taxonomic research combined with diligent and intelligent collecting but it is also partly due to the immigration or at least the increasing frequency of some species. Twenty- five species (ca 7% of the flora) have not been recorded during the present century, and may be extinct, but many more have declined in frequency. The paper seeks to elucidate the extent and the causes of such changes; for this purpose much evidence from other districts is reviewed.

Apart from the physical destruction of local specialized habitats and the creation of new habitats, the more important causes of widespread change have been: (a) eutrophication of both land and water, partly as a result of (b) changes in agricultural practice, including the heavy application of fertilizers, and immediate ploughing and resowing after harvest; (c) a decline in grazing, especially by rabbits; (d) changes in forestry practice, including the cessation of coppicing and the planting of conifers, increasing the amount of decaying woody material; (e) most far-reaching of all, widespread but slight atmospheric pollution, the effect of which seems to be due to ‘acid rain’ rather than to toxic SO₂.

Factors (d) and (e), though adverse for basiphil species, have favoured acidiphil species, at least eight of which have increased in frequency. An additional four acidiphil species first recorded in the county since 1935 have remained rare, but evidence from other districts suggests that they are increasing.

Some dioecious mosses formerly fruited more freely in Oxfordshire than they do now.     

A Floristic Study of Flora of Seed Plants of the Chaqia-GongheBasin and Its Contiguous Zone in Qinghai,China

The Chaqia-Gonghe Basin and its contiguous zone is situated in eastern Qinghai of China, between latitude 34°45′- 37°00′N and longitude 98°45′- 101°30′E . Ranging from 2 800m to 5 305 m in altitude, the total area is about 38 300km2 . Its climate is continental or plateau-continental one . There are 854 species of native seed plants belonging to 59 families and 277 genera there , which occupied 37. 37% of the total species, 54 . 53% of the total genera and 65. 56% of the total families in Qinghai respectively . The floristic characteristics of the native seed plants are as follows : ( 1) Number of species and woody ones there are poor . (2) At generic level, the flora is temperate in nature, dominated by north temperate elements especially the typical elements from the warm and cold zone of Eurasia , but also with some elements of warm, cold warm and alpine ones . (3 ) The area is a margin of distribution for many species or genera especially for tropical ones, and thus , the flora of the area is clearly marginal in nature . (4) Floristic-geographically , it is a converged and transitional region for the Qinghai-Tibetan Plateau Alpine Flora , Loess Plateau Warm Flora , as well as East Asia Alpine Desert Flora. (5 ) It is a part of the Tangute Flora . (6 ) It is a key area to divide the subregions in the flora of Qinghai on the Tangute Area .     

青海茶卡- 共和盆地及其毗邻地区种子植物区系

茶卡- 共和盆地及其毗邻地区位于青海省的东部, 约处于北纬34°45′～ 37°00′, 东经98°45′～ 101°30′。海拔2 800～5 305 m, 面积约38 300 km2 , 属于高原大陆性气候类型。本区共有野生种子植物59 科、277 属、854 种。分别占青海省总科数的65 . 56%、总属数的54 .53%、总种数的37 . 37%。区系特征如下:1、种类相对较贫乏, 木本更贫乏。2、就属的层面而言, 本区属于以北温带成分, 特别是欧亚大陆温、寒地带典型成分为优势的, 兼具旱生温性、耐寒温性和高寒类型的温带区系性质。3、许多种、属和热带类型均以本区为边缘分布区, 因而区系的边缘性质明显。4、为青藏高原高寒区系、黄土高原温性( 草原)植物区系和亚洲东部荒漠区系在我国的交汇过渡区, 区系的过渡性质明显。5、是青藏高原植物亚区唐古特地区的一部分。6、是青海植物区系或“唐古特植物地区”划分植物亚地区的关键区域。     

Species richness and endemicity in the Spanish vascular flora

Data from an updated and revised checklist of the Spanish flora is analyzed. The Spanish vascular plant flora is composed of 204 families, 1433 genera and 7071 species. Floristic data are analysed by considering three regions: mainland Spain with 5984 species, the Balearic Islands with 1521 species and Canary Islands with 2066 species. Extinct species are included in the analysis, with an extinction rate of 0.35% of the flora. A total of 1488 endemic species are recognized, which account for 21% of the Spanish flora. The rate of endemism in the Canary Islands is 25.9%, considerably higher than for the Balearic Islands (6.9%) and for mainland Spain (13.8%). A list of the 35 strict endemic genera is compiled, of which 65.7% are Canarian, and another list of 27 subendemic genera of which 48.1% are also Canarian. An estimated 12% of the Spanish flora is non‐native, with large variation between the 20.7% of the Canary flora, 10.5% of mainland Spain and 9.7% of the Balearic Islands. Finally, the composition of the Spanish flora for large groups and families is ana lysed. The data show that species richness and endemicity rates have been overestimated by previous authors, and are similar to other Mediterranean countries. It is concluded that a complete revision of the Macaronesian flora is strongly needed to obtain an accurate comparison with the flora of other areas.     

Origin and evolution of endemic plants of the Bonin (Ogasawara) Islands

The Bonin Islands are typical oceanic islands, located at the western part of the North Pacific Ocean and approximately 1,000 km south of mainland Japan. This archipelago consists of about 20 small islands. Although floristic diversity is low due to the small area and limited environmental diversity, the Bonin Islands harbor unique endemic flora as in other well-known oceanic islands. This paper presents a brief summary of the results obtained from recent studies on the endemic flora of the Bonin Islands. The results are reviewed in relation to the four stages of the evolution of endemic flora in the oceanic islands; migration, establishment, enlargement and diversification. The ancestors of the flora originated mostly from tropical and subtropical Southeast Asia or mainland Japan by rare events of long distance dispersal. The proportion of bird-dispersed species is relatively high as for other oceanic islands. Genetic data sets obtained from allozyme variation in some endemic species suggest that migration occurred several million years ago and genetic diversity is correlated with current population size. At the time of establishment, self-compatible plants are expected to have an advantage. However, the percentage of dioecious plants is relatively high. This is partly due to evolutionary changes from hermaphroditic ancestors to dioecy which occurred in two genera in the Bonin Islands. In addition, there are some examples of evolutionary changes from herbaceous ancestors to woody endemics. Adaptive radiation is found in some genera, although the number of congeneric endemic species is less than five. Studies of allozyme variation inPittosporum, Symplocos andCrepidiastrum showed that genetic identity is generally very high between congeneric species in spite of their distinct morphologies. This result suggests that divergence of these species occurred rather recently and distinct morphological differences are based on a limited number of genetic changes.     

Hybridization and introgression in a mixed population of the intertidal seaweeds Fucus evanescens and F. serratus

The introduced Fucus evanescens (hermaphroditic) and the native F. serratus (dioecious) have been in secondary contact along the Danish coast of the Kattegat Sea for 60–100 years and dioecious hybrids have been observed at Blushøj for several years. Hybridization in Fucus is unusual because it appears to always involve a hermaphroditic and a dioecious parental pair. We determined the degree and spatial patterns of introgression for 286 individuals using 10 microsatellite loci and cpDNA. Hybrids accounted for nearly 13% of the population, yet parental species were well differentiated (F_ST = 0.633). The presence of F. evanescens chloroplasts in 100% of F₁ hybrids revealed asymmetrical hybridization. Fucus evanescens cpDNA was observed in 50% of introgressed and 5.4% of pure F. serratus, but no F. serratus cpDNA was found in F. evanescens. In contrast, nuclear DNA introgression was symmetrical with an equal amount (≈1.5%) of genes introgressed into each parental species. Survivorship and viability data suggest selection against hybrids in the hybrid zone.     

Plant Invasions on an Oceanic Archipelago

Plant invasions are particularly noticeable on oceanic islands. For many ecological or evolutionary phenomena, oceanic islands offer advantages in comparison to continental regions, because they are often simpler systems. The Juan Fernández (Robinson Crusoe) Islands, located 667 km west of continental Chile, provide an especially favorable case study of plant invasions on an oceanic archipelago. They have little flora, no influence from aboriginal peoples, and good historical and recent documentation of flora, vegetation and human impacts. The total flora of the archipelago consists of 441 vascular plants, of which 209 are native (125 of them endemic) and 232 are aliens. Many alien species exist on the Juan Fernández Archipelago, but three shrubs are particularly invasive: Aristotelia chilensis, Rubus ulmifolius, and Ugni molinae. About 15% of the total area of the island has been impacted by at least one of these shrubs. In addition, the herbaceous Acaena argentea, now occurs at high abundance in 12% of the total area of the island. Comparisons of earlier and recent surveys of vegetation reveal that the area impacted by Aristotelia, Rubus, and Ugni molinae has increased tremendously. Among the most frequent aliens are Euro-Mediterranean taxa, also present on other archipelagos. A few aliens that are serious invasives on other archipelagos have been recently observed near San Juan Bautisata, the only village on the island. Effective measures involving stronger monitoring and sanitation of incoming materials (foodstuffs, building materials, etc.), cutting or poisoning of invasives, and reseeding with native species, are necessary to help preserve the native and endemic flora (and biota) of these islands.     

The taxonomy and ecology of the vascular plant flora of Singapore: a statistical analysis

A total of 2277 species of vascular plant in 868 genera and 158 families are recorded as native to the Republic of Singapore. This flora consists of 174 pteridophytes (7.6% of the total), eight gymnosperms (0.4%), 590 monocots (25.9%) and 1505 dicots (66.1%).Median values of species per family, genera per family and species per genus are 5, 2 and 1, respectively. The most abundant 'families' and genera in terms of species are Orchidaceae (196 spp.), Pteridophyta (174 spp.), Rubiaceae (140 spp.), Euphorbiaceae (116 spp.), Gramineae (100 spp.) and Eugenia (46 spp.), Ficus (43 spp.), Dendrobium (33 spp.), Bulbophyllum (29 spp.) and Cyperus (29 spp.). Trees constitute 37.2% of the flora, shrubs 9.8%, herbs 25.9%, climbers 14.0% and epiphytes 13%.Euphorbiaceae and Rubiaceae dominate the woody flora; pteridophytes, Gramineae and Cyperaceae the herbs; Rubiaceae, Palmae and Annonaceae the climbers and Orchidaceae and pteridophytes the epiphyteS. Species confined to coastal habitats comprise 11.1% of the total flora; 73.5% are lowland forest species and the remaining 15.5% are non-tree species of open and frequently-disturbed sites. The vascular plant of flora of Bukit Timah Nature Reserve, the largest remaining patch of primary rain forest in Singapore, is also analysed. Its flora is similar to that of the forest component of the complete Singapore flora. The Singapore flora, in comparison with other tropical floras, is highly diverse, but appears depauperate in epiphytes and climbers.     

A survey of floral traits, breeding systems, floral visitors, and pollination systems of the angiosperms of the Juan Fernández Islands (Chile)

A survey of the reproductive features of the angiosperm flora of the Juan Fernández Archipelago (Chile) is presented to provide a species-based review of reproduction and pollination, to identify generalizations associated with these systems, to understand the evolution of these features, and to utilize these data to promote conservation. The collection of original data was extensive, based on our own fieldwork, and was combined with data from existing literature. Data recorded include habit, sexual system, flower size, shape, and color, and the hypothesized pollination system of the first colonizers. In addition, the data on compatibility, presence and type of dichogamy, observed floral visitors, presence of floral rewards, and currently known pollination systems are summarized. Pairwise comparisons of different features are tested for statistical association. The flora is typically composed of perennials. The majority of the species have very small or small flowers. Inconspicuous flowers (i.e., a shape character describing flowers with no optical attraction) are widespread, as are dish-shaped flowers. Green is the most frequent flower color, followed by white and yellow. Most species are hermaphroditic, 9% are dioecious, and 9% are monoecious. Some 30% of the species are protandrous and 7% protogynous. Detailed studies of compatibility of about 14% of the flora indicate that 85% of these species are self-compatible (SC). Although most species studied are SC, their level of autogamy is low. Nevertheless, selfing mediated via geitonogamy is the most frequent mechanism of pollen transfer. Outcrossing is mainly achieved through dioecy and self-incompatibility (SI), promoted by dichogamy in the hermaphroditic flowers, and facilitated by wind pollination. About 55% of the species offer nectar rewards, and only 2% offer pollen rewards. Floral visitors are rare to uncommon. Two hummingbird species, one of them endemic, are considered as pollinators for 14 plant species. Flies, moths, and beetles are the native insect visitors to flowers, but they have been documented on only 11 plant species (7%). Even insect visits to these few species were rarely observed. Given the infrequent, irregular, and imprecise nature of native insect association with flowers, there is no certainty that any of the species are truly insect pollinated. Two species of introduced ants and a new endemic bee were recorded as well; however, neither is likely currently important to the pollination of the native flora species. About 9% of the extant flora is currently bird pollinated, and we hypothesize that 47% is wind pollinated. However, we propose that most of the colonizers were ancestrally either insect or wind pollinated. There is association between a number of current floral features and the hypothesized pollination of colonizers. Therefore, to a large extent the flower color, shape, and size of the extant flora may express the pollination syndromes of colonizers rather than representing extant pollination. In addition, the presence of nectar in many species of extant flora does not necessarily indicate biotic pollination. Thus, studies of the reproductive biology on oceanic island plants need to be conducted species by species before broad generalizations can be made, because the observed features can be misleading. Possible changes in the pollination system were assessed by comparison of species for which there are reliable data with the hypothesized pollination of their colonist progenitors. The wind-or bird-pollinated species have retained the pollination system of the colonizers. In other instances, species seem to bear a different pollination system: from ancestral insect systems to current hummingbird-or wind-pollination systems. The lack of alternative means of biotic pollination seems to have led in a number of instances to anemophily—in essence a default pollination system. The lack of strong selection pressure for wind pollination and the relative youth of the archipelago may help explain why the features associated with wind pollination in these species are not so obvious. Because there are many recorded extinctions of vascular plants from islands versus those from continental areas, it is imperative to invest additional effort in protecting the remaining island species. Conservation or restoration programs cannot be effective without a deep and broad understanding of the reproductive biology of the plants. In order to conserve these plants, programs must involve a combination of reproductive and environmental measures. The ultimate fate of some species may depend on preserving the plant-hummingbird relationship, including the web of organisms that affect both plant and pollinator. The populations of introduced animals and weeds must be controlled. Experimentally produced allogamous seeds would enhance diversity in restoration programs. In addition, the preservation of habitat seems to be the central challenge to indirectly protect the unique island species.     

Taxonomic and biogeographic patterns in the native and alien woody flora of Kashmir Himalaya,India

Biodiversity inventories unravel insightful taxonomic and biogeographic patterns that provide valuable inputs in guiding conservation practices and policymaking. Here we present an updated inventory of the native and alien woody flora of the Kashmir Himalaya, India, together with analyses of the patterns of taxonomic composition, geographic distribution, and invasion status of the alien species. The data was assembled from the floristic literature of the last two centuries, supplemented with herbarium records and validated by field surveys over the past decade. In total, the woody flora comprised of 520 species; out of these 322 species were native and 198 alien, yielding a proportion of 38% aliens. Amongst the natives, 16 species are on the IUCN red list 2009. While 37 genera included both native and alien species indicating a potential for hybrid formation, only 15 families were represented by alien species exclusively. Regarding the invasion status, presently 37 alien woody species are naturalized and 7 are invasive in the region. The results show that the Kashmir Himalaya harbours a diverse woody flora. It is concluded that a distinction between the native and the alien flora, as adopted in the present study, is crucial for a meaningful analysis of any biodiversity patterns. We predict that in the near future many alien woody species, presently naturalized in the region, can become serious plant invaders, and therefore need further research efforts and timely management.     

The vascular flora of Rome

The aim of this paper is to document plant diversity in the metropolitan area of Rome (Italy) by providing a comprehensive inventory of the present-day vascular flora and an overview of its composition and species diversity. We compiled the floristic catalogue by including all vascular plant entities that occur spontaneously within the administrative boundaries of the Rome municipality. The data, which were gathered from extensive field surveys, from a broad review of the literature and from herbaria records, were updated and integrated in a comprehensive account. The inventory of the flora of metropolitan Rome lists 1649 entities, from 139 families and 677 genera. The flora contains 228 taxa that are non-native to the Italian flora, 81 of which are established in the study area. The overview of flora reveals a remarkable species diversity and outlines the main characteristics of the flora of Rome, which may be summarized as (1) a large number of taxa of high conservation value which occur in remnants of near-natural vegetation, (2) the loss or decline of some species, particularly of native freshwater plants, (3) a remarkably high native species richness within the urban area, which includes many native woody species and (4) a rich ruderal flora, prevalently composed of native species that are well adapted to human disturbance, along with a variety of taxa of non-native or uncertain origin. The large set of data and the overview presented in this paper represent a fundamental framework for future research and for the conservation of plant diversity in metropolitan Rome.     

Sexual Systems and Ecological Correlates in an Azonal Tropical Forests, SW China

Unlike the seriate lowland rain forests in SE Asia, the tropical vegetation of Xishuangbanna (SW China) has developed in habitats with comparatively lower temperatures and precipitation. Consequently, although most of the families and genera of the flora are components of tropical ecosystems, many have reached their distribution limits in latitude. Selection pressures on sexual systems in these environments may be different from that experienced in lowland tropical regions. Here, we examine the sexual systems of 685 species of flowering plants belonging to 109 families and 356 genera based on 42 plots with a total area of 15.4 ha and compare our results with surveys of sexual systems from other tropical ecosystems. Among these species, 60.6 percent were hermaphroditic, 14.3 percent were monoecious, and 25.1 percent were dioecious. The percentage of dioecious sexual system among tree species (26.1%) was similar or higher than that of other tropical tree floras. Monoecy was significantly associated with the tree growth form and was relatively common in seasonal forest vegetation. Sexual systems involving unisexual flowers (dicliny) are particularly well represented in the tropical forests of Xishuangbanna accounting for 39.4 percent of all species surveyed. This pattern may be associated with the prevalence of relatively small inconspicuous flowers in the tropical forests of SE Asia and their correlation with diclinous sexual systems generally.     

Dioecy and its correlates in the flowering plants

Considerable effort has been spent documenting correlations between dioecy and various ecological and morphological traits for the purpose of testing hypotheses about conditions that favor dioecy. The data analyzed in these studies, with few exceptions, come from local floras, within which it was possible to contrast the subsets of dioecious and nondioecious taxa with regard to the traits in question. However, if there is a strong phylogenetic component to the presence or absence of dioecy, regional sampling may result in spurious associations. Here, we report results of a categorical multivariate analysis of the strengths of various associations of dioecy with other traits over all flowering plants. Families were scored for presence of absence of monoecy or dioecy, systematic position, numbers of species and genera, growth forms, modes of pollination and dispersal, geographic distribution, and trophic status. Seven percent of angiosperm genera (959 of 13,500) contain at least some dioecious species, and ≈6% of angiosperm species (14,620 of 240,000) are dioecious. The most consistent associations in the data set relate the presence of dioecy to monoecy, wind or water pollination, and climbing growth. At both the family and the genus level, insect pollination is underrepresented among dioecious plants. At the family level, a positive correlation between dioecy and woody growth results primarily from the association between dioecy and climbing growth (whether woody or herbaceous) because neither the tree nor the shrub growth forms alone are consistently correlated with a family's tendency to include dioecious members. Dioecy appears to have evolved most frequently via monoecy, perhaps through divergent adjustments of floral sex ratios between individual plants. Monoecy itself is related to abiotic pollination and climbing growth as revealed by multivariate analysis. Dioecy and monoecy are concentrated in the less advanced superorders of Thorne (1992) and subclasses of Cronquist (1988). The frequency of dioecy found in a local flora therefore reflects the level of dioecy in its particular pool of families as much as, or more than, local selective factors. The positive associations of dioecy with abiotic pollination and monoecy are related to floral developmental and morphological attributes, as is the negative association with bird and bat pollination; the positive association of dioecy with climbing growth is tentatively explained in terms of differential selection for optimal resource allocation to sexual function. If rapid upward growth is at a premium in climbers and if fruit set at least temporarily inhibits growth or requires the production of thicker, more slowly growing stems to support heavy fruits, it might be advantageous to postpone femaleness. If the effect is strong, this may favor male plants.     

Determinants of regional and local patterns in the floras of braided riverbeds in New Zealand

Aim To determine the composition and biogeographical origins of the native and naturalized flora of braided shingle riverbeds in New Zealand and whether the proportion of naturalized species is greater than in similar habitats elsewhere in the world. To test whether regional species pools, landscape matrix configuration, and local riverbed environment are all equally important in determining community structure in these systems. Location The braided reaches of four catchments on each of the eastern and western sides of North Island and South Island, New Zealand. Methods Plotless records of all native and naturalized seed plant and fern species on disturbed mobile surfaces were made over the length of four rivers. Altitude, climate variables, riverbed attributes, and surrounding land‐cover from maps were recorded for each site. For all species, the taxonomy, life form, origin and history, and presence within the province through which the rivers flowed, was obtained from published floras. Direct and indirect ordination and variance partitioning were employed to examine how native and naturalized species composition varies among and within rivers, and the degree to which this variation reflects climate, characteristic of riverbeds, and the surrounding land‐cover. Regression was used to determine how much introduction date and native geographical range influence the frequency of naturalized species. Results The total riverbed flora of 289 species comprises 40% native species and 60% naturalized species, both dominated by Asteraceae and Poaceae. The relative contributions of other plant families differ, and the two groups comprise different life‐form spectra. Native species occur across fewer rivers (mean 1.6 rivers) than naturalized species (mean 2.1 rivers). Species common in at least one river system tend to be widespread, occurring in at least three rivers. The rivers differ in their floras, with distinctions between North Island and South Island, and eastern and western rivers. The South Island rivers have more native species and a higher proportion of their regional native species pools than North Island rivers, whereas they have a lower proportion of their naturalized species pools. Introduction date and native geographical range are correlated with frequency of individual naturalized species. Geographical position and climate, riverbed variables of substrate size and seepage presence, and the type of adjacent land‐cover, differ between rivers and are significantly related to species composition. Land‐cover variables alone account for 46% of the explained variation in species composition, environmental variables alone 32% and components of the environment and land‐cover that co‐vary, 22%. Main conclusions New Zealand shingle riverbeds are among the riparian communities in the world most invaded by naturalized species but comparisons are difficult because studies of similar riverbeds are rare. New Zealand riverbeds are dominated by short‐lived terrestrial species from Eurasia. Native species are most frequent in South Island rivers draining extensive high mountains, and their abundances are determined to a greater extent by riverbed processes than are those of the naturalized species, which are more abundant when the surrounding landscape is modified. The distribution of the naturalized flora is not yet at equilibrium with the environment. Each river has a distinctive flora determined by ambient environmental factors, aspects of the riverbed environment itself, adjacent land‐cover reflecting the presence of native vegetation, the intensity of human modification or use, and invasion history.