Weeds as crops: The value of maize field weeds in the valley of Toluca, Mexico

Maize field weeds or agrestals are widely used in central Mexico as potherbs (quelites) and forage. This work presents quantitative data on these uses from the village of San Bartolo del Llano, Municipio de Ixtlahuaca, Valley of Toluca, an area with a relatively intensive, semicommercial agriculture. We interviewed 24 families of the village and 10 vendors at the market of Ixtlahuaca regularly during one rainy season (1995) on type and quantity of weed use. Also, the weed vegetation was surveyed and we interviewed 49 farmers on their farming practices and on costs. All of the 74 weed species found in maize fields were useful as forage, potherb, medicinal, or ornamental. Within the village, 11 species were eaten; an average family consumed 4.5 kg of wild potherbs per month during the rainy season. In Ixtlahuaca, 2150 kg of 10 species were sold, worth 3054 pesos (US $611). For quantity and gross economic value, forage was much more important. On the average, 1 ha of maize field produced a harvest of 1.5 t of green forage, worth about 25% of the gross value of the maize harvest, and 55% of its net value. The combination of maize with forage weeds for stabled animals constitutes an interesting integrated farming system. The weeds increase the useful biomass of the field, improve nutrition of the farmers, do not reduce the yield of the main crop, as the fields are kept weed free during the critical period, and provide erosion control, shade, and green manure.     

Dispersal failure contributes to plant losses in NW Europe

The ongoing decline of many plant species in Northwest Europe indicates that traditional conservation measures to improve the habitat quality, although useful, are not enough to halt diversity losses. Using recent databases, we show for the first time that differences between species in adaptations to various dispersal vectors, in combination with changes in the availability of these vectors, contribute significantly to explaining losses in plant diversity in Northwest Europe in the 20th century. Species with water- or fur-assisted dispersal are over-represented among declining species, while others (wind- or bird-assisted dispersal) are under-represented. Our analysis indicates that the 'colonization deficit' due to a degraded dispersal infrastructure is no less important in explaining plant diversity losses than the more commonly accepted effect of eutrophication and associated niche-based processes. Our findings call for measures that aim to restore the dispersal infrastructure across entire regions and that go beyond current conservation practices.     

Human-mediated dispersal of seeds over long distances

Human activities have fundamental impacts on the distribution of species through altered land use, but also directly by dispersal of propagules. Rare long-distance dispersal events have a disproportionate importance for the spread of species including invasions. While it is widely accepted that humans may act as vectors of long-distance dispersal, there are few studies that quantify this process. We studied in detail a mechanism of human-mediated dispersal (HMD). For two plant species we measured, over a wide range of distances, how many seeds are carried by humans on shoes. While over half of the seeds fell off within 5m, seeds were regularly still attached to shoes after 5 km. Semi-mechanistic models were fitted, and these suggested that long-distance dispersal on shoes is facilitated by decreasing seed detachment probability with distance. Mechanistic modelling showed that the primary vector, wind, was less important as an agent of long-distance dispersal, dispersing seeds less than 250 m. Full dispersal kernels were derived by combining the models for primary dispersal by wind and secondary dispersal by humans. These suggest that walking humans can disperse seeds to very long distances, up to at least 10 km, and provide some of the first quantified dispersal kernels for HMD.     

Evolution of black yeasts: possible adaptation to the human host

Ascomycetous black yeasts show adaptations to a wide array of environmental conditions. Dothideaceous black yeasts are mostly found on plant leaves, while among herpotrichiellaceous species there are numerous opportunists on humans.Factors which are of ecological significance include the presence of melanin and carotene, formation of thick cell walls and meristematic growth, presence of yeast-like phases, presence of additional forms of conidiogenesis, thermo- and osmotolerance, adhesion, hydrophobicity, production of extracellular polysaccharides, siderophores and acidic or alkaline secondary metabolites.The potential pathogenicity of a species is partly determined by its natural ecological niche. Dothideaceous black yeasts are osmotolerant rather than pathogenic. Herpotrichiellaceous black yeasts probably have low competitive ability and are found in rather special niches as secondary saprophytes, e.g., on bacterial mats, on other fungi or in poor environments. Some species possibly utilize animal vectors for dispersal.     

Ramesh Gulati: a personal approach,based on his retirement lecture

Historical and modern migrations and dispersal of most marine organisms (intertidal, benthic, meiofaunal, planktonic, nektonic, or neustonic) are classically interpreted in terms of their natural dispersal potential. Exceptions are introduced species, largely recognized since the 19th century, known to have been transported by human activities. However, humans were transporting species along coastlines and across oceans for millennia and centuries prior to the advent of the first biological surveys. Thus, the presumptive natural distributions of many species may be questioned. Reviewed here are some basic concepts about invasions of non-native species. Human activities move species isolated in time and space from other oceans or continents, and thus human-mediated transport does not simply speed up natural dispersal processes. Both past and modern-day invasions are often overlooked, leading to an underestimation of the scale of invasion diversity and impact. Because vectors, donor regions, and recipient regions change over time, invasions will continue along long-standing but un-managed corridors. The impact of most invasions has never been studied and, therefore, it is not possible to conclude that most invasions have no impact, nor is it generally possible to say that invasions have become `integrated' into a community or ecosystem in ecological time. Finally, invasions in the ocean are not limited to harbours and ports, but are found in a wide variety of marine habitats, ranging from the open ocean continental shelf to exposed rocky shores. The existence of human-mediated vectors has created extraordinary challenges to our understanding and interpretation of the ecology, biogeography, evolutionary biology, and conservation biology of marine communities.     

A reassessment of the role of simple seed traits in mortality following herbivore ingestion

Grazing mammals are regarded as major vectors in seed dispersal of grassland plants, through seed ingestion and subsequent excretion (endozoochory). The (evolutionary) ecology underlying this dispersal mode is relatively poorly understood because there are limited data, among others, on how seed attributes perform in this process (and could thus be selected for). For seed mortality following ingestion, contrasting patterns described for the role of simple seed traits seem partly due to inadequate comparative analytical methods. We conducted a feeding experiment in which controlled seed quantities from 48 grassland herb and grass species were fed to cattle. Seed mass, length and shape measurements were related to seed mortality rate using phylogenetically independent contrasts, which account for taxonomic interdependence. The proportion of seeds surviving the digestive tract was generally low, but it reached 100%, with increased germination for some species. Neither seed size nor shape correlated significantly with mortality. Structural traits are likely to overrule these simple seed traits, as illustrated by high survival of species having water-impermeable seeds. As this trait has interesting properties with respect to intraspecific variation and genetics, it could provide opportunities for inference on evolutionary consequences for this type of dispersal.     

Scale-dependence and mechanisms of dispersal in freshwater zooplankton

Communities of organisms form as a result of both interspecific and abiotic interactions within local habitat patches and dispersal among patches in a region. Local processes are expected to play a dominant role when dispersal occurs much more often than extinction. We performed two field experiments to examine rates and mechanisms of dispersal in freshwater pond zooplankton communities. First, we tested the effect of distance from a source on the rate of colonization of artificial habitat by placing wading pools at 5, 10, 30 and 60 m from two natural fishless ponds and observing the succession of zooplankton. Seventy-eight percent of the species in the source ponds that were capable of living in the pools colonized at least once during the experiment. A new species was found in the pools on average once every four days, suggesting that colonization events occur on the order of days to weeks for many species. Colonization rates declined further from the source at one pond but not the other, and the effect of distance was relatively weak at both ponds. This suggests that many species disperse broadly over short distances. The second experiment tested the role of animal vectors for zooplankton dispersal by restricting access to the pools. Eight treatments were imposed that excluded potential animal vectors along a body size gradient from large mammals to small insects. While the treatments affected zooplankton colonization, many species invaded even when all animals larger than 1 mm were excluded. Animal vectors may therefore be less important for dispersal than wind. Our results suggest that zooplankton are highly effective dispersers over short distances, and can disperse via several mechanisms. Local interactions should therefore play a dominant role in structuring these communities at small regional scales.     

Picky hitch‐hikers: vector choice leads to directed dispersal and fat‐tailed kernels in a passively dispersing mite

Dispersal is a central life‐history trait for most animals and plants: it allows to colonize new habitats, escape from competition or avoid inbreeding. Yet, not all species are mobile enough to perform sufficient dispersal. Such passive dispersers may use more mobile animals as dispersal vectors. If multiple potential vectors are available, an active choice can allow to optimize the dispersal process and to determine the distribution of dispersal distances, i.e. an optimal dispersal kernel. We explore dispersal and vector choice in the neotropical flower mite Spadiseius calyptrogynae using a dual approach which combines experiments with an individual‐based simulation model. Spadiseius calyptrogynae is found in lowland rainforests in Costa Rica. It inhabits inflorescences of the understorey palm Calyptrogyne ghiesbreghtiana and is phoretic on a number of flower visitors including bats, beetles and stingless bees. We hypothesised that the mites should optimise their dispersal kernel by actively choosing a specific mix of potential phoretic vectors. In a simple olfactometer setup we showed that the flower mites do indeed discriminate between potential vectors. Subsequently we used an individual‐based model to analyse the evolutionary forces responsible for the observed patterns of vector choice. The mites combine vectors exhibiting long‐distance dispersal with those allowing for more localized dispersal. This results in a fat‐tailed dispersal kernel that guarantees the occasional colonization of new host plant patches (long distance) while optimizing the exploitation of clumped resources (local dispersal). Additionally, kin competition results in a preference for small vectors that transport only few individuals at a time. At the same time, these vectors lead to directed dispersal towards suitable habitat, which increases the stability of this very specialized interaction. Our findings can be applied to other phoretic systems but also to vector‐based seed dispersal, for example.     

Does seed dispersal limit initiation of primary succession in desert playas?

To investigate the initiation of primary succession in a cold-desert playa-dune complex, we studied the large-scale (2000 m) seed (diaspore) dispersal patterns at Mono Lake, California. Seeds of seven of the ten species reaching the barren playa had wind-dispersal adaptations. Rates of dispersal (numbers of seeds per square metre per day) were as much as three orders of magnitude lower on the playa than in the diverse dune vegetation. However, seed input appeared sufficient to reach potential safe sites on the playa, with a peak input of 66 ± 8 total seeds·m·d. The smooth playa surface, the virtual absence of aboveground barriers, and the high windspeed environment promote the long-distance dispersal of seeds (at least 1300 m for Chrysothamnus spp. and at least 700 m for Sarcobatus vermiculatus). The large spatial scale of sampling revealed a relatively high seed input onto the playa by the dominant pioneer species S. vermiculatus, despite the low abundance of parent vegetation in this region. All of these results implicate low rates of seed entrapment as an obstacle to establishment on this desert playa, rather than a lack of seed input.     

SEED DISPERSAL OF TRILLIUM OVATUM (LILIACEAE) IN SECOND-GROWTH REDWOOD FORESTS

Trillium ovatum (Liliaceae) is myrmecochorous: its seeds bear large elaiosomes that are attractive to ants. Nevertheless, in coastal second-growth redwood forests of northern California, most seedlings occur in mixed-age clusters close to potential parents, suggesting that seed dispersal is limited. Ants were absent or rare at two relatively cool, moist study sites. At these sites, most seeds either eventually fell passively from fruits or were knocked to the ground by banana slugs that foraged on the elaiosomes. At two warmer, drier sites, a single species of ant, Lasius pallitarsis, dispersed the seeds but tended to remove the elaiosomes before returning to the nest. Thus at all sites a large number of seeds remained close to adults, accounting for the observed pattern of seedling distribution. The dispersal adaptations of T. ovatum and other redwood forest myrmecochores probably evolved in forests where seed-carrying ants were more common.     

Does seed retention potential affect the distribution of plant species in highly fragmented calcareous grasslands?

Long-distance dispersal is a crucial factor in the life-cycle of plants, especially in our modern, highly fragmented landscapes. Because natural herds of large animals have disappeared and grazing practices have been abandoned, important potential vectors for seed dispersal over large distances may have been lost. In the context of the re-establishment of grazing management for nature conservation purposes, it is therefore important to gain insight in the ability of grazing animals to act as seed dispersal vectors. Whereas local dispersal mainly occurs through standard vectors typically described based on morphological adaptations of the diaspore, large herbivores act as non-standard seed dispersers. Therefore, traditional dispersal classes are loosing scientific relevance and continuous predictors of dispersal potential have been proposed. Here, we explored whether dispersal related plant traits, including the "seed retention potential", could explain the distribution patterns of 180 plant species over 64 fragmented semi-natural calcareous grasslands in Belgium. The distribution of habitat specialist plant species was strongly determined by the degree of isolation of the grasslands. Interestingly, species distribution patterns were clearly linked with a species' potential to migrate through large grazers, as quantified by its retention potential: species producing seeds with high retention capacity were less affected by habitat isolation. Categorical dispersal classes based on seed morphology did not explain a species' response to fragment isolation. Although seed retention potential outperformed simple seed dimensional traits, plant height, which is an indicator of epizoochorous attachment potential, was even more important. Therefore we suggest further extension of the epizoochorous retention potential model by incorporating basic ecological mechanisms that effectively contribute to successful dispersal events.     

A general framework for propagule dispersal in mangroves

Dispersal allows species to shift their distributions in response to changing climate conditions. As a result, dispersal is considered a key process contributing to a species' long‐term persistence. For many passive dispersers, fluid dynamics of wind and water fuel these movements and different species have developed remarkable adaptations for utilizing this energy to reach and colonize suitable habitats. The seafaring propagules (fruits and seeds) of mangroves represent an excellent example of such passive dispersal. Mangroves are halophytic woody plants that grow in the intertidal zones along tropical and subtropical shorelines and produce hydrochorous propagules with high dispersal potential. This results in exceptionally large coastal ranges across vast expanses of ocean and allows species to shift geographically and track the conditions to which they are adapted. This is particularly relevant given the challenges presented by rapid sea‐level rise, higher frequency and intensity of storms, and changes in regional precipitation and temperature regimes. However, despite its importance, the underlying drivers of mangrove dispersal have typically been studied in isolation, and a conceptual synthesis of mangrove oceanic dispersal across spatial scales is lacking. Here, we review current knowledge on mangrove propagule dispersal across the various stages of the dispersal process. Using a general framework, we outline the mechanisms and ecological processes that are known to modulate the spatial patterns of mangrove dispersal. We show that important dispersal factors remain understudied and that adequate empirical data on the determinants of dispersal are missing for most mangrove species. This review particularly aims to provide a baseline for developing future research agendas and field campaigns, filling current knowledge gaps and increasing our understanding of the processes that shape global mangrove distributions.     

Tree dispersal strategies in the littoral forest of Sainte Luce (SE-Madagascar)

Zoochory is the most common mode of seed dispersal for the majority of plant species in the tropics. Based on the assumption of tight plant-animal interactions several hypotheses have been developed to investigate the origin of life history traits of plant diaspores and their dispersers, such as species-specific co-evolution, the low/high investment model (low investment in single fruits but massive fruiting to attract many different frugivores versus high investment in single fruits and fruit production for extended periods to provide food for few frugivores), and the evolution of syndromes which represent plant adaptations to disperser groups (e.g. birds, mammals, mixed). To test these hypotheses the dispersal strategies of 34 tree species were determined in the littoral forest of Sainte Luce (SE-Madagascar) with the help of fruit traps and tree watches. The impact of fruit consumers on the seeds was determined based on detailed behavioral observations. Phenological, morphological and biochemical fruit traits from tree species were measured to look for co-variation with different types of dispersal. No indication for species-specific co-evolution could be found nor any support for the low/high investment model. However dispersal syndromes could be distinguished as diaspores dispersed by birds, mammals or both groups (mixed) differ in the size of their fruits and seeds, fruit shape, and seed number, but not in biochemical traits. Five large-seeded tree species seem to depend critically on the largest lemur, Eulemur fulvus collaris, for seed dispersal. However, this does not represent a case of tight species-specific co-evolution. Rather it seems to be the consequence of the extinction of the larger frugivorous birds and lemurs which might also have fed on these large fruits. Nevertheless these interactions are of crucial importance to conserve the integrity of the forest.     

Report on wind dispersal in a lowland moist forest in central French Guiana

An analysis of the flowering plant flora of a lowland moist forest in central French Guiana reveals 298 species with adaptations for wind dispersal. This represents 16.2% of the flowering plant flora and 9.8% of the class Magnoliopsida (dicotyledons). The most diverse wind-dispersed families are the Orchidaceae in the Liliopsida (monocotyledons) with 135 species and the Bignoniaceae in the Magnoliopsida with 37 species. The wind-dispersed species of central French Guiana have evolved either small, dust-like seeds, fruits or seeds with various kinds of wings, fruits or seeds with tufts of hairs, or expanded wing- or parachute-like persistent calyces. Most wind-dispersed species, among the liliopsids, are epiphytes and, among the magnoliopsids, trees or lianas. In central French Guiana, collections of these species with mature diaspores have been gathered most often in October and March, the months with peak wind velocities. In contrast, collections from June and July, when wind velocities are at a minimum, are rare.     

Emus as non-standard seed dispersers and their potential for long-distance dispersal

Long-distance seed dispersal may have important consequences for species range, migration rates, metapopulation dynamics, and gene flow. Plants have evolved various adaptations for seed dispersal by standard agents, with typical dispersal distances associated with them. Seeds may also be dispersed by non-standard agents for which they do not show any apparent adaptation and may reach long distances. By sampling the droppings of emus Dromaius novaehollandiae at three localities in Western Australia, we investigated their potential to act as long-distance dispersers of seeds with adaptations for dispersal modes other than endozoochory, such as unassisted, ant, wind and exozoochory, for which they act as non-standard agents. Seventy-seven plant species with five types of dispersal syndromes were found in the 112 droppings analysed, with at least 68 having viable seeds. Although endozoochory was the most frequent syndrome, the presence of other syndromes was important in terms of number of species (61%) and seeds (50%). Estimates of species richness indicated that an increase in sampling effort would increase the number of species observed, especially among non-endozoochores. As a consequence of their long gut retention times and high mobility, emus can provide long-distance dispersal opportunities that may be especially relevant for species with dispersal modes of typically short distances (unassisted, ant).
Our results suggest that the role of emus as non-standard agents for long-distance dispersal should be taken into account for understanding current geographic ranges, gene flow and metapopulation dynamics of some plant species, as well as for predicting their future responses to climate change and fragmentation.     

How do invasive species travel to and through urban environments?

Globalisation has resulted in the movement of organisms outside their natural range, often with negative ecological and economic consequences. As cities are hubs of anthropogenic activities, with both highly transformed and disturbed environments, these areas are often the first point of entry for alien species. We compiled a global database of cities with more than one million inhabitants that data had on alien species occurrence. We then identified the most prominent pathways of introduction and vectors of spread of alien species in these cities. Most species were intentionally introduced to cities and were released or escaped from confinement. The majority of alien species then spread within cities through natural means (primarily unaided dispersal). Pathway prominence varied across the taxonomic groups of alien species: the most prominent pathway for plants and vertebrates was the escape pathway; for invertebrates the stowaway and contaminant pathways were most likely to facilitate introductions. For some organisms, pathway prominence varied with the geographical and climatic characteristics of the city. The characteristics of the cities also influenced the prominence of vectors of spread for alien species. Preventing the natural spread of alien species within cities, and into adjacent natural environments will be, at best, difficult. To prevent invasions, both the intentional and unintentional introduction of potentially harmful alien species to cities must be prevented. The pathways of introduction and vectors of spread identified here should be prioritised for management.     

Interactions between propagule pressure and seed traits shape human-mediated seed dispersal along roads

Human-mediated dispersal along the road network is a crucial process in the population dynamics of roadside vegetation and during plant invasions. The potential for a species to be dispersed by vehicles is, however, difficult to quantify. The predictive power of categorical classification schemes of human-mediated dispersal is limited as many species that are usually attributed to particular primary dispersal vectors may become subject to very different secondary dispersal vectors owing to human activity. Analysing seed traits that promote seed transport by human dispersal vectors could overcome these limitations. However, the analysis has to account for the divergent chance of seed transport that results from different propagule pressures along the transport corridor.To reveal the effects of seed traits and their interplay with propagule pressure on the chance and magnitude of human-mediated dispersal by vehicles, we compared traits and regional frequencies of a set of species that were dispersed by vehicles to a control set not dispersed but present in the same study area. We then used the same traits for a comparison of intentionally and unintentionally introduced species with the flora of Berlin.Different traits influenced the chance of vehicle dispersal and its magnitude. While propagule pressure was most important for determining the magnitude of seed transport, small seed mass and size best predicted the absolute chance of species dispersal by vehicles. The dispersal of nonnative species was least dependent on propagule pressure.Seed traits that were important in vehicle dispersal were similarly reflected in unintentionally introduced species in the Berlin flora. Mean seed size of these species was lower compared to the entire Berlin flora, whereas it was higher for intentionally introduced species. This suggests that unintentional introduction of nonnative plant species pre-selects for seed traits that promote further spread by human-mediated adhesive dispersal.Probability and magnitude of adhesive seed transport by vehicles can be predicted by dispersal-related plant traits. However, the effect size of plant traits on dispersal strongly depends on regional propagule pressure. This highlights the need to analyse interactions between species traits and propagule pressure.     

Nonsignificant isolation by distance implies limited dispersal

Understanding the scale of dispersal is an important consideration in the conservation and management of many species. However, in species in which the high‐dispersal stage is characterized by tiny gametes or offspring, it may be difficult to estimate dispersal directly. This is the case for many marine species, whose pelagic larvae are dispersed by ocean currents by several days or weeks before beginning a benthic, more sedentary, adult stage. As consequence of the high‐dispersal larval stage, many marine species have low genetic structure on large spatial scales (Waples 1998 ; Hellberg 2007 ). Despite the high capacity for dispersal, some tagging studies have found that a surprising number of larvae recruit into the population they were released from (self‐recruitment). However, estimates of self‐recruitment are not informative about mean dispersal between subpopulations. To what extent are limited dispersal estimates from tagging studies compatible with high potential for dispersal and low genetic structure? In this issue, a study on five species of coral reef fish used isolation by distance (IBD) between individuals to estimate mean dispersal distances (Puebla et al. 2012 ). They found that mean dispersal was unexpectedly small (<50 km), given relatively low IBD slopes and long pelagic durations. This study demonstrates how low genetic structure is compatible with limited dispersal in marine species. A comprehensive understanding of dispersal in marine species will involve integrating methods that estimate dispersal over different spatial and temporal scales. Genomic data may increase power to resolve these issues but must be applied carefully to this question.     

Epizoochorous dispersal by barbs, hooks, and spines in a lowland moist forest in central French Guiana

An analysis of the flowering plant flora of a lowland moist forest in central French Guiana reveals 25 species with adaptations for epizoochorous dispersal by barbs, hooks, or spines (stick-tights). This represents 1.5% of the flowering plant flora. Stick-tights are represented among the monocotyledons of the flora of central French Guiana by five species of Poaceae. The Fabaceae and Asteraceae, each with six species with stick-tight dispersal, are the richest families of flowering plants in terms of stick-tight dispersal in central French Guiana. This region possesses numerous potential mammal and bird dispersal agents, therefore it is a puzzle why stick-tight dispersal, especially among species of the canopy, is not more frequent.     

Larval dispersal of the invasive fall armyworm,Spodoptera frugiperda,the exotic stemborer Chilo partellus,and indigenous maize stemborers in Africa

Larval dispersal either through ballooning or crawling results in a redistribution of the insect population and infestations within and between plants. In addition, invasive species, such as the fall armyworm (FAW), Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae), and the exotic stemborer Chilo partellus (Swinhoe) (Lepidoptera: Crambidae), may displace indigenous stemborers on maize in Africa. To test whether larval dispersal activity may play a role in the displacement of indigenous stemborers, larval dispersal was compared between FAW, C. partellus, and the indigenous species Busseola fusca (Fuller) and Sesamia calamistis (Hampson) (both Lepidoptera: Noctuidae). Twenty potted maize plants were infested with one batch of eggs either from stemborers (B. fusca, S. calamistis, or C. partellus) or from FAW and monitored in the greenhouse for ballooning activities. After egg hatching, both ballooning and non-ballooning larvae were identified according to species and counted. FAW neonate larvae had greater potential for ballooning off than stemborers, irrespective of species. For each species, more females dispersed than males, and their survival rate was higher than that of non-ballooning larvae. In addition, plant-to-plant larval movements were studied using 6.25-m² plots of caged maize in a completely randomized design with five replicates. FAW was found to have wider dispersal and plant damage potential than any of the stemborer species. In conclusion, in contrast to C. partellus, the invasive characteristic of FAW can be explained, in part, by its higher larval dispersal activity compared to stemborers. This difference in larval dispersal might also be considered in sampling plans for monitoring pest density in the field.