Integration of invasive <Emphasis Type="Italic">Opuntia</Emphasis> spp. by native and alien seed dispersers in the Mediterranean area and the Canary Islands

The success of many alien plant species depends on mutualistic relationships with other species. We describe the assemblage of seed dispersers on three species of alien Opuntia invading Mediterranean and Macaronesian habitats, and examine the quality of such plant-animal interactions. We identified vertebrates consuming O. maxima, O. dillenii and O. stricta fruits by direct observation and collecting droppings and pellets. Phenology of the alien species, as well as that of coexisting native species, was monitored for an entire year. Germination tests of ingested and non-ingested seeds were performed both in the greenhouse and in the field. Seed coat thickness and viability were also measured for all treatments. A great variety of taxa, including reptiles, birds and mammals actively participate in the seed dispersal of Opuntia. Phenology of Opuntia fruits in Menorca and Tenerife overlaps with only a few native fleshy-fruited plants present in the study areas, which suggests an advantage for the invader. Most seeds germinated during the second year of the experiment, independently of the effect produced by the dispersers’ guts. We found great variation in the germination percentage of Opuntia after gut passage and in the effects of ingestion on seed coat thickness. Seed viability was somewhat reduced after gut passage compared to manually depulped seeds. Our results show how different Opuntia species are integrated into native communities by means of mutualistic interactions, with both native and alien dispersers. Although with heterogeneous effects, either type of disperser potentially contributes to the spread of these alien cacti in the recipient areas.     

The role of fruit diet within a temperate breeding bird community in southern Spain

In southern temperate regions many fleshy-fruited plant species ripen their fruits during the breeding or early post-breeding season and these are consumed by resident birds. I studied the frugivore community of the summer-fruiting tree Frangula alnus ssp. baetica in southern Spain and the significance of fruit in the diet of adult and recently fledged birds. Twelve species consumed F. alnus fruits. Almost 97% of the foraging visits were carried out by the four most abundant species, Robin Erithacus rubecula, Blackcap Sylvia atricapilla, Blackbird Turdus merula and Blue Tit Parus caeruleus. During the ripening season Robins and Blackcaps concentrated at fruit-rich sites. Entire families of Blackcap and Blackbird regularly entered fruiting trees together. In contrast, adult Robins always foraged solitarily, and accounted for only 4% of the tree visits, while 96% were realized by recently fledged birds of the first brood. In all species, young birds foraged as efficiently as adults. F. alnus fruits are not part of the regular nestling diet, but they are a significant part of the diet for recently fledged Robins during their postnatal dispersal. Conversely, these are the most important and effective seed dispersers while adult Robins scarcely take part in the dispersal of F. alnus seeds.     

Non-redundancy in seed dispersal and germination by native and introduced frugivorous birds: implications of invasive bird impact on native plant communities

Seed dispersal by vertebrate animals is important for the establishment of many fleshy-fruited plant species. Different frugivorous species can provide different seed dispersal services according to their specific dietary preferences as well as behaviour and body traits (e.g. body size and beak size of birds). Our aim was to study redundancies and complementarities in seed dispersal and germination between the two main native seed disperser birds and the introduced silver pheasant Lophura nycthemera in the temperate Patagonian forests. For this, we collected fresh droppings from the studied species and analyzed seed content. We conducted germination trials for four plant species common in bird droppings; two native species (Aristotelia chilensis and Rhaphithamnus spinosus) and two invasive non-native species (Rubus ulmifolius and Rosa rubiginosa). Both native frugivorous birds and the silver pheasant dispersed fruits of non- native fleshy-fruited plants, but their roles were non-redundant in terms of species dispersed and effect on seed germination. The silver pheasant dispersed a proportionally high number of non-native seeds, while native birds dispersed a high number of native seeds. In addition, the effect of gut treatment in seed germination differed between seed dispersers. Native birds promoted the germination for the two native plant species studied, while the silver pheasant promoted the germination of one non-native plant. This suggests that seed dispersal by the silver pheasant may contribute to the spread of some invasive fleshy-fruited plants in the ecosystems that otherwise would not be dispersed by any other bird. The understanding of redundancies and complementarities on seed dispersal and germination between native and introduced birds will allow improving the management of fleshy-fruited non-native plants.     

Different contributions of birds and mammals to seed dispersal of a fleshy-fruited tree

Birds and mammals are important seed dispersers of fleshy-fruited plants. Although their behaviors are different, they frequently consume the same species. Thus to understand the dispersal of fleshy-fruited plants, the contribution of birds and mammals to seed dispersal has to be evaluated. Besides, within birds or mammals, some species may functionally different from others. In this study, we examined seed dispersal of the fleshy-fruited tree Swida controversa focusing on the difference between two frugivore groups (birds and mammals), and differences between species within groups. Collected seeds and S. controversa trees were identified using simple sequence repeat (SSR) genotyping, thus enabling to determine the distance between mother tree and dispersed seeds. The avian species were identified by DNA barcoding of feces, whereas the mammalian species were identified by the shape and smell of feces. Most seeds that fell near or under the maternal trees were dispersed by birds, resulting in short seed dispersal distances (average, 13 m). DNA barcoding detected five taxa of avian dispersers. No differences were detected in seed dispersal distance by different avian taxa (i.e., the distance between dispersed seeds and their maternal trees within the research plot); however the rate of seed immigration from outside the research plot by some avian taxa varied significantly. The seed dispersal distance by mammals was significantly further (127 m; min > 50 m) than that by birds. Additionally, immigrated seeds accounted for approximately two-thirds of mammal-dispersed seeds, indicating that these seeds were from outside the research plot, and that mammals significantly contributed to the long-distance seed dispersal of S. controversa. No differences in seed dispersal distance were detected between different mammalian taxa. Overall, this study revealed that birds and mammals show clearly different seed dispersal patterns, and thus, they play different roles in the regeneration of S. controversa.     

The role of avian frugivores in germination of seeds of fleshy-fruited invasive alien plants

Many highly invasive plant species have fleshy fruits which are eaten by native frugivorous animals. These frugivores play an important role in long-distance seed dispersal, and may also affect germination success. The aim of this study was to determine whether generalist frugivores enhance or decrease seed germination of invasive alien species through pulp removal or seed coat abrasion, besides serving as dispersal agents. Fruits of four fleshy-fruited invasive alien plant species, namely Solanum mauritianum, Cinnamomum camphora, Lantana camara and Psidium guajava, were fed to three generalist avian frugivorous species, which have been observed feeding on these fruits in the wild. Seed retention time was recorded as this affects dispersal distance and the duration that seeds are exposed to the effects of the gut. Seeds removed from excreta, seeds from manually de-pulped fruit, and whole fruit were planted in soil trays housed in a greenhouse. Daily germination counts were done. Seed retention times differed significantly between bird species for all fruits, except those of C. camphora. However, all frugivores had a similar effect on the germination success of seeds of S. mauritianum, L. camara and P. guajava, showing that gut retention time was not important. Germination of seeds from manually de-pulped fruits did not differ from that of ingested seeds of all plant species, suggesting that seed coat abrasion was also not important. Pulp removal resulted in significantly higher germination rates, both in the two species with larger, multi-seeded fruit (S. mauritianum and P. guajava), and in the two species having single-seeded fruit with waxy exocarps (C. camphora and L. camara). Pulp removal also resulted in significantly earlier germination of L. camara and P. guajava seeds. Therefore, frugivores not only accelerate dispersal, but also greatly enhance seed germination of all fleshy-fruited invasive alien species in this study.     

Could poor seed dispersal contribute to predation by introduced rodents in a Hawaiian dry forest?

Dry forests are among the most diverse, yet threatened, communities in Hawai’i. Dry forests throughout the archipelago suffer from a lack of natural regeneration of trees. Two factors that may limit tree recruitment include poor seed dispersal and seed predation by rodents. Poor or limited dispersal of fleshy-fruited species results in seeds and fruits falling directly under parents. Dispersed and non-dispersed seeds may differ in their vulnerability to predation. We tested effects of seed location (under/away from parent trees) and pulp (presence/absence) on predation of four native species that suffer from limited dispersal and one readily-dispersed alien species in Kanaio Natural Area Reserve, Maui. Three natives (Diospyros sandwicensis, Pleomele auwahiensis, Santalum ellipticum), had significantly more seeds removed under parent trees than in exposed sites away from trees. For the one alien (Bocconia frutescens) and two native trees (D. sandwicensis, P. auwahiensis) that were evaluated, significantly more intact fruits were removed than were cleaned seeds. Presence of teeth marks and gnawed seed husk fragments indicate introduced rodents are destroying many of the seeds they remove. These results suggest that seed predation is disproportionately concentrated among poorly-dispersed seeds and may contribute to recruitment failure.     

Impact of avian frugivores on dispersal and recruitment of the invasive <Emphasis Type="Italic">Prunus serotina</Emphasis> in an agricultural landscape

Although seed dispersal is considered to be a key process determining the spatial structure and spread of non-native plant populations, few studies have explicitly addressed the link between dispersal vector behaviour, seed distribution and seedling recruitment to gain insight into the process of exotic species invasion within a fragmented landscape context. The present study analyses the relationship between avian frugivory and spatial patterns of seed deposition and seedling recruitment for an expanding population of the invasive Prunus serotina in a hedgerow network landscape in Flanders, Belgium. We quantified fruit production, observed frugivores, and determined the spatial distribution of bird droppings and P. serotina seedlings. A relatively diverse assemblage of frugivores visited P. serotina seed trees, with Columba palumbus and Turdus merula being by far the most important dispersers. Landscape structure strongly affected dispersal vector behaviour and the spatial distribution of perching birds, droppings and seedlings. Frugivorous birds non-randomly dispersed seeds to perching sites and an association between perching birds, seed deposition and seedling recruitment was found. Results indicate that landscape structure contributes to non-random seed deposition of P. serotina by common local frugivores. Cutting the larger seed trees is proposed as the most feasible measure to slow down the invasion rate.     

The role of avian ‘seed predators’ as seed dispersers

Seed dispersal is a central process in plant ecology with consequences for species composition and habitat structure. Some bird species are known to disperse the seeds they ingest, whereas others, termed ‘seed predators’, digest them and apparently play no part in dispersal, but it is not clear if these are discrete strategies or simply the ends of a continuum. We assessed dispersal effectiveness by combining analysis of faecal samples and bird density. The droppings of seed dispersers contained more entire seeds than those of typical seed predators, but over a quarter of the droppings of seed predators contained whole seeds. This effect was further magnified when bird density was taken into account, and was driven largely by one frequent interaction: the Chaffinch Fringilla coelebs, a typical seed predator and the most abundant bird species in the area and dispersed seeds of Leycesteria formosa, a non‐native plant with berry‐like fruits. These results suggest the existence of a continuum between seed predators and seed dispersers.     

Wahlberg’s epauletted fruit bat (<Emphasis Type="Italic">Epomophorus wahlbergi</Emphasis>) as a potential dispersal agent for fleshy-fruited invasive alien plants: effects of handling behaviour on seed germination

The spread of invasive alien plants into natural habitats is of growing global concern. Several studies have investigated the role that avian frugivores play in the dispersal of these seeds and their effects on germination success. Fruit bats have however received little attention as important dispersal agents of invasive alien plants, despite their recognized role as long distance dispersal agents of various native flora. We investigated whether Wahlberg’s epauletted fruit bats, Epomophorus wahlbergi, would positively influence the germination of seeds of invasive alien plants. These fruit bats were fed fruits of four invasive alien plant species—Psidium guajava, Melia azedarach, Eriobotrya japonica, and Morus alba. Epomophorus wahlbergi were able to process more fruit per gram body mass than birds have been observed to do. Spat and de-pulped control seeds had similar germination success and germinated at approximately the same time for most species. While seeds retained in whole fruit had significantly less germination success than spat seeds for all species, except M. azedarach, they mostly germinated at approximately the same time. Epomophorus wahlbergi can swallow small seeds (< 2 mm), while seeds larger than this are generally spat out. Large fruit are usually carried away to feeding roosts where seeds are dropped, thereby dispersing seeds and fruits which are too large for some bird species to ingest. Epomophorus wahlbergi should not be underestimated as dispersers of these invasive alien plants as they consume proportionally large amounts (0.62 ± 0.09 to 0.99 ± 0.11 g.g⁻¹ body mass) of fruit, except for M. azedarach, and positively affect their seed germination rates.     

Spatial and temporal patterns of rowan (<Emphasis Type="Italic">Sorbus aucuparia</Emphasis> L.) regeneration in West Carpathian subalpine spruce forest

Non-random seed shadows are commonly seen in plant species whose seeds are dispersed by animals, in particular by birds. The behaviour of birds can influence the spatial pattern of seed dispersal and, consequently, the entire regeneration process of fleshy-fruited trees. This study examined regeneration patterns in a fleshy-fruited tree species, rowan (Sorbus aucuparia L.), growing in West Carpathian subalpine spruce forests, focussing on two problems: the temporal relationship between rowan regeneration and gap formation, and the spatial relationship between rowan regeneration and stand structure. It was found that rowan seedlings and saplings were recruited in advance of gap formation. Establishment of new rowan individuals in gaps was infrequent, but gaps enhanced their regeneration nearby under spruce canopy, where they occurred densely in a narrow belt about 15 m wide. Inside spruce stands, the highest density of young rowans was directly under crowns, especially near trunk bases. Few rowan saplings were found growing under mature rowan trees. The presence of a rowan seedling and sapling bank determines whether rowans fill spruce stand gaps. Dense rowan groves can develop mainly in extensive but slowly expanding gaps.     

Patterns of Seed Dispersal and Dispersal Failure in a Hawaiian Dry Forest Having Only Introduced Birds

Dry forests are among the most endangered natural communities in the Hawaiian Islands. Most have been reduced to isolated trees and small forest fragments in which native tree species reproduce poorly. The replacement of native birds by introduced generalists may be contributing to dry forest decline through modification of seed dispersal patterns. To document seed dispersal by introduced birds, we conducted foraging observations on fleshy-fruited trees and measured seed rain under trees and in adjacent open areas for 1 year in a dry forest dominated by native trees. Although trees covered only 15.2 percent of the study area, 96.9 percent of the bird-dispersed seeds were deposited beneath them. The Japanese white-eye (Zosterops japonicus) was the principal dispersal agent. Among bird-dispersed seeds, those of the invasive tree Bocconia frutescens accounted for 75 percent of all seeds collected beneath trees (14.8 seeds/m²/yr) and the invasive shrub Lantana camara accounted for 17 percent. Although nearly 60 percent of the reserve's native woody species possess fleshy fruits, introduced birds rarely disperse their seeds. Native trees accounted for <8 percent of all bird-dispersed seeds and are consequently experiencing dispersal failure by falling directly under parent trees. Smaller-seeded non-native plants, in contrast, may be benefiting from dispersal by introduced birds. Current dispersal patterns suggest that these readily disseminated non-native plants may eventually replace the remaining native flora.     

The potential role of waterbirds in dispersing invertebrates and plants in arid Australia

1. The role of waterbirds as vectors of plants and invertebrates within and between arid‐zone wetlands is poorly understood. We present the first detailed study of passive dispersal by nomadic birds in Australasia. We investigated the numbers and types of invertebrate and plant propagules within freshly collected faecal samples as well as their viability. We compared dispersal among Grey Teal (Anas gracilis), Eurasian Coot (Fulica atra) and Black Swan (Cygnus atratus) in the Macquarie Marshes, a complex of temporary to semi‐permanent wetlands in New South Wales. 2. When faecal samples (n = 60) were inundated in the laboratory and monitored over 3 weeks, ciliates (75% of samples), nematodes (22%), ostracods (13%) and rotifers (5%) were recorded, with higher taxon richness in coot samples. Faecal samples (n = 71) were also sieved to quantify intact propagules, and ostracod eggs (70% of samples), large branchiopod eggs (31%) and bryozoan statoblasts (31%) were the most abundant invertebrates. Diaspores of 19 plant taxa were recorded, 14 of which were germinated in the laboratory or shown to be viable at the end of germination trials. The abundance and diversity of invertebrate propagules was highest in coot samples, whereas the abundance and diversity of diaspores was highest in teal samples. 3. One Australian Pelican (Pelecanus conspicillatus) sample was obtained and found to contain more taxa and far more propagules than any sample from other waterbirds, suggesting that piscivorous birds might have an important role in the indirect dispersal of propagules ingested by fish. 4. Our results support a role for birds in explaining the distributions of cosmopolitan plant genera such as Lemna, Typha, Myriophyllum and Nitella. The alien plants Ranunculus sceleratus, Medicago polymorpha and Polygonum arenastrum were recorded, demonstrating the potential role of waterfowl in the spread of exotic species. As the frequency and duration of flooding of arid‐zone wetlands decreases owing to human activities, the importance of waterbirds in facilitating recolonisation of temporary wetlands is likely to increase.     

Generalized avian dispersal syndrome contributes to Chinese tallow tree (Sapium sebiferum, Euphorbiaceae) invasiveness

Abstract. Plants possessing generalized dispersal syndromes are likely to be more invasive than those relying on specialist dispersal agents. To address this issue on a local and regional scale, avian seed dispersal of the invasive alien Chinese tallow tree (Sapium sebiferum (L.) Roxb.) was assessed in forests and spoil areas of South Carolina and along forest edges in Louisiana during the 1997–99 fruiting seasons. Tallow trees in these floristically distinct habitats had a few common and many casual visitors, and considerable species overlap among habitats was found. However, bird species differed in the importance of dispersing and dropping seeds among habitats. Important dispersal agents common to forests and spoil areas of South Carolina included Northern Flicker, American Robin and Red‐winged Blackbird, whereas Red‐bellied Woodpecker and European Starling were important in the former and latter habitat, respectively. In Louisiana, Red‐bellied Woodpecker, American Robin, Northern Cardinal and Eastern Bluebird dispersed many seeds. Nearly all species foraging on seeds were winter residents. Estimated numbers of seeds dispersed and dropped were higher in spoil areas of South Carolina than in Louisiana because of higher numbers of individuals per visit, higher seed consumption and seed dropping rates, and longer foraging durations. Within South Carolina, more seeds were dispersed and dropped in spoil areas than in forests because of higher numbers of birds per visit. These findings show that among habitats, tallow tree attracts diverse but variable coteries of dispersal agents that are qualitatively similar in seed usage patterns. We suggest that its generalized dispersal syndrome contributes to effective seed dispersal by many bird species throughout its range. Effects of differential avian use among locales may include changes in local bird communities, and differing tallow tree demographics and invasion patterns.     

Shorebirds as important vectors for plant dispersal in Europe

Shorebirds (Charadriiformes) undergo rapid migrations with potential for long‐distance dispersal (LDD) of plants. We studied the frequency of endozoochory by shorebirds in different parts of Europe covering a broad latitudinal range and different seasons. We assessed whether plants dispersed conformed to morphological dispersal syndromes. A total of 409 excreta samples (271 faeces and 138 pellets) were collected from redshank Tringa totanus, black‐winged stilt Himantopus himantopus, pied avocet Recurvirostra avosetta, northern lapwing Vanellus vanellus, Eurasian curlew Numenius arquata and black‐tailed godwit Limosa limosa in south‐west Spain, north‐west England, southern Ireland and Iceland in 2005 and 2016, and intact seeds were extracted and identified. Godwits were sampled just before or after migratory movements between England and Iceland. The germinability of seeds was tested. Intact diaspores were recovered from all bird species and study areas, and were present in 13% of samples overall. Thirteen plant families were represented, including Charophyceae and 26 angiosperm taxa. Only four species had an ‘endozoochory syndrome’. Four alien species were recorded. Ellenberg values classified three species as aquatic and 20 as terrestrial. Overall, 89% of seeds were from terrestrial plants, and 11% from aquatic plants. Average seed length was higher in redshank pellets than in their faeces. Six species were germinated, none of which had an endozoochory syndrome. Seeds were recorded during spring and autumn migration. Plant species recorded have broad latitudinal ranges consistent with LDD via shorebirds. Crucially, morphological syndromes do not adequately predict LDD potential, and more empirical work is required to identify which plants are dispersed by shorebirds. Incorporating endozoochory by shorebirds and other migratory waterbirds into plant distribution models would allow us to better understand the natural processes that facilitated colonization of oceanic islands, or to improve predictions of how plants will respond to climate change, or how alien species spread.     

Crayfish invasion facilitates dispersal of plants and invertebrates by gulls

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Differential seed dispersal systems of endemic junipers in two oceanic Macaronesian archipelagos: the influence of biogeographic and biological characteristics

This article evaluates the seed dispersal systems of two congeneric and endemic fleshy-fruited plants in the context of two relatively close oceanic archipelagos. For this purpose, representative populations of the endangered junipers Juniperus cedrus in the Canary Islands and Madeira, and Juniperus brevifolia in the Azores were studied. Despite both species sharing the same biogeographic region, we set out to test whether different conditions of the islands and biological characteristics of each juniper species determine the distinctive guilds of seed dispersers involved. We assessed the quantitative and qualitative role of the potential frugivores, showing that the wintering Turdus torquatus and the native Turdus merula were the main seed dispersers for J. cedrus and J. brevifolia, respectively (Frequency of occurrence: 74.9%, 80.2%; germination increase with respect to controls: 11.6%, 15.5%; for J. cedrus and J. brevifolia, respectively). The endemic lizard Gallotia galloti was quantitatively outstanding as seed disperser of J. cedrus, although its qualitative effect does not appear to be beneficial. The introduced rabbit Oryctolagus cuniculus acts as a disruptor in both natural seed dispersal systems, as inferred from the high percentage of damaged seeds found in their droppings. Our results indicate that J. cedrus and J. brevifolia are primarily adapted to ornithochory processes, T. torquatus and T. merula being their respective legitimate long-distance dispersers. Although these birds should be playing a key role in the connectivity of fragmented populations, the dependence of J. cedrus on a migrant bird involves a notable fragility of the system.     

The role of migratory ducks in the long‐distance dispersal of native plants and the spread of exotic plants in Europe

Little is known about the role of migratory waterfowl in the long‐distance dispersal (LDD) of seeds. We studied the gut contents of 42 teals Anas crecca collected in the Camargue, southern France, and found intact seeds of 16 species. There was no relationship between the probability that a given seed species was found intact in the lower gut, and the seed hardness or size. The number of seeds found in the oesophagus and gizzard (a measure of ingestion rate) was the only significant predictor of the occurrence of intact seeds in the lower gut, so studies of waterfowl diet can be used as surrogates of dispersal potential. In a literature review, we identified 223 seed species recorded in 25 diet studies of teal, pintail Anas acuta, wigeon A. penelope or mallard A. platyrhynchos in Europe. We considered whether limited species distribution reduces the chances that a seed can be carried to suitable habitat following LDD. Overall, 72% of plant species recorded in duck diets in southern Europe (36 of 50) were also recorded in the north, whereas 97% of species recorded in duck diets in the north (137 of 141) were also recorded in the south. This suggests a great potential for LDD, since most dispersed plants species occur throughout the migratory range of ducks. Migratory ducks are important vectors for both terrestrial and aquatic plant species, even those lacking the fleshy fruits or hooks typically used to identify seeds dispersed by birds. Finally, we show ducks are important vectors of exotic plant species. We identified 14 alien to Europe and 44 native to Europe but introduced to some European countries whose seeds have been recorded in duck diet.     

Herb endozoochory by cockatoos: Is ‘foliage the fruit’?

Establishing whether herb seed endozoochory is accidental or has evolved independently or in combination with other dispersal mechanisms may be valuable in the study of plant–animal interactions, but it remains unexplored for birds. We tested whether an Australian cockatoo, the galah (Eolophus roseicapilla), swallows entire seeds when feeding on other tissues without subsequent seed digestion, thus enhancing seed dispersal (the ‘foliage is the fruit’ hypothesis). Our preliminary sampling provides strong evidence supporting that this seed predator also acts as a legitimate endozoochorous disperser. A large proportion of droppings contained numerous seeds of six herb species of three plant families, surviving gut passage to be dispersed as viable propagules. The wide range in the number of seeds found in combinations with up to five species in particular droppings suggests both simultaneous and sequential passive ingestion without seed digestion and/or focused seed predation and digestion. As expected for inadvertent ingestion and inefficient digestion, our findings suggest that seed number and richness of dispersed plants are associated traits in this particular mutualistic interaction. This relationship can have important implications in community‐wide processes, favouring herbs whose seeds are disseminated in a viable state over those predated or negatively affected by gut transit.     

Birds as key vectors for the dispersal of some alien species: Further thoughts

W. Solarz, K. Najberek, A. Pociecha & E. Wilk‐Wo?niak ( 2017 , Diversity and Distributions, 23 , 113–117) published a letter in Diversity and Distributions debating our view that waterbirds are important vectors of alien species (C. Reynolds, N. A. F. Miranda & G. S. Cumming, 2015 Diversity and Distributions, 21 , 744–754; A. J. Green, 2016 Diversity and Distributions, 22 , 239–247) and question whether future research into the mechanisms under‐pinning this phenomenon can be advantageous for the practical management of alien species. Additionally, Solarz et al. suggest that human activities are the primary source of all alien species introductions and that waterbirds may only act as vectors of secondary dispersal. In this letter, we respond to several arguments raised by the authors surrounding the relevance of waterbird‐mediated dispersal in the introduction and spread of alien species. We emphasize the partly deterministic nature of waterbird dispersal and the significance of long‐distance dispersal events (and hence the potential for primary introductions of new alien species across political boundaries). Finally, we reaffirm the importance of further research into dispersal by birds to improve our capacity to foresee and manage invasions of those alien species with strong capacity to spread via avian vectors.     

Seasonally different modes of seed dispersal in the prostrate annual, <Emphasis Type="Italic">Chamaesyce maculata</Emphasis> (L.) Small (Euphorbiaceae), with multiple overlapping generations

The modes of seed dispersal in the prostrate annual, Chamaesyce maculata, with multiple overlapping generations were investigated. We found that C. maculata has two modes of seed dispersal; autochory in the summer and myrmecochory in the autumn. Seasonally different modes of seed dispersal have not been known in other plant species. The large proportion of seeds produced in the summer was positioned further than the expanse of the parent plants by automatic mechanical seed dispersal. Therefore, autochory would be effective for avoiding competition between parent and offspring plants. No autochory occurred in the seeds produced in the autumn. The seeds of C. maculata without an elaiosome were dispersed by seed-collecting ants in the autumn. Although 18 ant species in total visited the plants of C. maculata at the 50 sites investigated, only two ant species, Tetramorium tsushimae and Pheidole noda frequently carried the seeds of C. maculata. The low frequency of seeds carried out of the nest by P. noda suggests that the workers of P. noda carry the seeds as food into their nest. So, P. noda might be a less effective seed disperser for C. maculata, corresponding to the effectiveness of seed dispersal by harvester ants. However, T. tsushimae ants frequently carried the seeds into and out of their nest, suggesting that T. tsushimae do not regard the seeds of C. maculata as a food resource. Thus, T. tsushimae may be an effective seed disperser for C. maculata.