Time-scale effects in the interaction between a large and a small herbivore

In the short term, grazing will mainly affect plant biomass and forage quality. However, grazing can affect plant species composition by accelerating or retarding succession at longer time-scales. Few studies concerning interactions among herbivores have taken the change in plant species composition into account. In a salt-marsh system, the long-term effects of exclusion of a large herbivore (cattle) on the abundance of a small herbivore (hare) were studied. Excluding cattle grazing for 30 years resulted in large changes in vegetation composition. In general, the cover of tall-growing species increased in the absence of cattle grazing. These long-term changes negatively affected hare grazing intensity. Hares preferentially fed on Festuca rubra and negatively selected tall growing plants, such as Elymus athericus, both in cattle-grazed and long-term ungrazed areas. However, the intensity of hare grazing was not related to the cover of F. rubra. The cover of tall-growing plants (E. athericus, Atriplex prostrata and Juncus maritimus) appeared to be the best predictor and hare grazing intensity decreased sharply with an increase of the cover of tall plants. When cover of tall plants did not increase, hare grazing intensity was not affected. The study shows that the time-scale of the experiment is of prime importance in studying interactions between herbivores. Species that do not seem to influence the abundance of one another or are competing for the same resources on a short time-scale might well be facilitating each other when looking at larger time-scales while taking plant species replacement into account.     

Vegetation,microclimate and soils associated with the latest-lying snowpatches in Australia

Background : The Snowy Mountains contain Australia's longest-lasting snowpatches. Because of climate change, their longevity has declined, with the loss of some specialist vegetation in the underlying snowbeds.

Aims: To characterise the current status of the vegetation associated with the longest-lasting snowpatches in Australia and its association with abiotic factors.

Methods: We assessed plant composition, soil depth, moisture and nutrients and subsurface temperatures in five zones of increasing vegetation height and cover in snowbeds.

Results: The zone beneath the middle of snowpatches was characterised by little vegetation cover and lower species richness, later emergence from snow, skeletal soils, and lower mean soil temperatures than zones further downslope where soils increased in depth and nutrient levels. Vegetation beneath these snowpatches no longer occurs in distinct communities. Plants have not simply migrated upslope, instead, areas that have deep soil that used to have snowpatch specialist species are being colonised upslope by grasses and downslope by tall alpine herbfield species that prefer bare ground.

Conclusions: Reduced longevity of Australia's longest-lasting snowpatches has led to the loss of distinct snowpatch plant communities. With limited soils beneath the centre of current snowpatches, and a lack of other suitable sites there is no location for these plant communities to migrate to.     

The impact of shrub browsing by mountain hare and reindeer in subarctic Sweden

Background: Climate warming has been causing an increase in tall shrub cover around the Arctic, however, mammalian herbivory has been shown to inhibit shrub expansion. Though the effect of reindeer (Rangifer tarandus) and many other mammals has been widely studied in this context, the role of the mountain hare (Lepus timidus) in subarctic Scandinavia remains unknown.

Aims: To quantify browsing from mountain hare and reindeer on tall shrubs in different vegetation types and to investigate differences in shrub preference between the two.

Methods: In the summers of 2013 and 2014, we counted signs of browsing by hare and reindeer on tall shrub species in 31 study plots at three alpine locations in the Scandes range, Sweden.

Results: Hare browsing was significantly more frequent than that by reindeer in two (dry-mesic heath and dry meadow) out of seven vegetation types studied. Reindeer browsing was significantly higher in the low herb meadow and Långfjället shrub heath. Two shrub species, Betula nana and Salix hastata, were significantly more browsed by hare, while reindeer browsing was significantly higher on S. phylicifolia and S. lapponum.

Conclusions: Our results show that mountain hares can cause extensive damage to tall shrubs in the subarctic and may have a stronger impact on shrub communities than previously recognised.     

Synergistic influences of introduced herbivores and fire on vegetation change in northern Patagonia,Argentina

Question: We investigated how cattle and European hares, the two most widespread exotic herbivores in Patagonia, affect species composition, life‐form composition and community structure during the first 6 years of vegetation recovery following severe burning of fire‐resistant subalpine forests and fire‐prone tall shrublands. We asked how the effects of introduced herbivores on post‐fire plant community attributes affect flammability of the vegetation. Location: Nahuel Huapi National Park, northwest Patagonia, Argentina Methods: We installed fenced plots to exclude livestock and European hares from severely burned subalpine forests of Nothofagus pumilio and adjacent tall shrublands of N. antarctica. The former is an obligate seed reproducer, whereas the latter and all other woody dominants of the shrubland vigorously resprout after burning. Results: Repeated measures ANOVA of annual measurements over the 2001‐2006 period indicate that cattle and hare exclusion had significant but complex effects on the cover of graminoids, forbs, climber species and woody species in the two burned community types. Significant interactions between the effects of cattle and hares varied by plant life forms between the two communities, which implies that their synergistic effects are community dependent. Conclusions: Following severe fires, the combined effects of cattle and hares inhibit forest recovery and favour transition to shrublands dominated by resprouting woody species. This herbivore‐induced trend in vegetation structure is consistent with the hypothesis that the effects of exotic herbivores at recently burned sites contribute to an increase in the overall flammability of the Patagonian landscape.     

Charismatic microfauna alter cyanobacterial production through a trophic cascade

The trophic ecology of cyanobacterial blooms is poorly understood on coral reefs. Blooms of toxic cyanobacteria, Lyngbya majuscula, can quickly form large mats. The herbivorous sea hare, Stylocheilus striatus, and the predatory nudibranch, Gymnodoris ceylonica, often associate with these blooms, forming a linear food chain: nudibranch—sea hare—cyanobacteria. Using laboratory studies, this study quantified (1) the functional response of nudibranchs, (2) the effect of sea hare size on predation rates, and (3) the strength of the indirect effect of sea hare predation on cyanobacteria (i.e., a trophic cascade). Nudibranchs consumed on average 2.4 sea hares d^?1, with the consumption of small sea hares 22 times greater than the consumption of large sea hares. Predation of sea hares reduced herbivory. Cyanobacterial biomass was 1.5 times greater when nudibranchs were present relative to when nudibranchs were absent. Although sea hare grazing can substantially reduce cyanobacterial biomass, predation of sea hares may mitigate grazing pressure, and therefore increase the abundance of cyanobacteria.     

Notes on the Scottish flora

Background: The degradation of alpine meadows on the Qinghai-Tibetan Plateau (QTP) has an impact on vegetation recruitment from seedlings and ramets.

Aims: Understanding the relative contribution of recruitment by seedling and ramet in alpine meadows is for the ecological restoration of degraded grasslands on the QTP.

Methods: An experiment was conducted to investigate seedling and ramet densities, species composition and their relationships with standing vegetation in plots representative of non-degraded (ND), lightly (LD), moderately (MD) and severely degraded (SD) alpine meadows.

Results: With increasing degradation and the reduction in vegetation cover, the number of seedlings and ramets declined. The proportion of young plants arising from seedlings was low, with the majority of species reproducing clonally. The establishment of seedlings of forbs in SD meadows led to species-rich, forb-dominant vegetation in these areas.

Conclusions: LD and MD meadows appear to be able to be managed by reduced grazing for the restoration of grass- and sedge-dominated pastures. In contrast, restoration of SD meadows will require additional intervention, such as of seeding and weed eradication.     

Multiscale habitat relationships of snowshoe hares (Lepus americanus) in the mixed conifer landscape of the Northern Rockies,USA: Cross‐scale effects of horizontal cover with implications for forest management

Snowshoe hares (Lepus americanus) are an ecologically important herbivore because they modify vegetation through browsing and serve as a prey resource for multiple predators. We implemented a multiscale approach to characterize habitat relationships for snowshoe hares across the mixed conifer landscape of the northern Rocky Mountains, USA. Our objectives were to (1) assess the relationship between horizontal cover and snowshoe hares, (2) estimate how forest metrics vary across the gradient of snowshoe hare use and horizontal cover, and (3) model and map snowshoe hare occupancy and intensity of use. Results indicated that both occupancy and intensity of use by snowshoe hares increased with horizontal cover and that the effect became stronger as intensity of use increased. This underscores the importance of dense horizontal cover to achieve high use, and likely density, of snowshoe hares. Forest structure in areas with high snowshoe hare use and horizontal cover was characterized as multistoried with dense canopy cover and medium‐sized trees (e.g., 12.7–24.4 cm). The abundance of lodgepole pine (Pinus contorta) was associated with snowshoe hare use within a mixed conifer context, and the only species to increase in abundance with horizontal cover was Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa). Our landscape‐level modeling produced similar patterns in that we observed a positive effect of lodgepole pine and horizontal cover on both occupancy and use by snowshoe hares, but we also observed a positive yet parabolic effect of snow depth on snowshoe hare occupancy. This work is among the first to characterize the multiscale habitat relationships of snowshoe hares across a mixed conifer landscape as well as to map their occupancy and intensity of use. Moreover, our results provide stand‐ and landscape‐level insights that directly relate to management agencies, which aids in conservation efforts of snowshoe hares and their associated predators.     

Competition between two grass species with and without grazing over a productivity gradient

Soil nutrient-level and herbivory are predicted to have opposing effects on the allocation pattern of the competitive dominant plant species. Lower stem and higher leaf allocation are favoured when plants are grazed, whereas a higher stem allocation is favoured at high nutrient levels. Grazing by hares and geese can prevent invasion of the tall Elymus athericus, into short vegetation of Festuca rubra, at unproductive stages of salt-marsh succession but not at more productive stages. We hypothesise that the negative effect of herbivory on Elymus decreases due to increasing soil nitrogen levels and shifts the competitive balance towards this species. We tested how simulated grazing and nitrogen availability affected the competitive balance between adult plants of both grass species in a greenhouse experiment. Elymus had a higher above-ground biomass production, invested relatively more in stem and root tissue and had a larger shoot length than Festuca. The above-ground relative yield of Elymus in mixtures of both species increased with increasing nitrogen levels. This indicates that Elymus was the superior competitor at high soil fertility. Although clipping removed relatively more biomass from Elymus than from Festuca and exceeded the observed biomass removal in field conditions, it did not change the competitive balance between both species. Decreasing effects of herbivory due to increasing nitrogen levels are not a likely explanation for the invasion of Elymus in productive marshes. The results suggest that once Elymus has established it can easily invade vegetation dominated by Festuca irrespective of grazing by herbivores such as hares and geese. Herbivory by small herbivores may mainly retard the invasion of this plant by influencing establishment itself.     

Interactions between hare and brent goose in a salt marsh system; evidence for food competition?

In this study we accumulate evidence that brown hare competes with brent goose for food resources in a temperate salt marsh. We show that both species overlap in habitat use and share food plants. The two herbivores mainly used the common habitat at different times of the day, with hares active in the dark and geese during the daylight. During the morning and evening, however, the habitat was exploited simultaneously. Food availability was manipulated by excluding brent geese on both small-scale (30 m²) and large-scale (0.96 ha) plots, while hares had free access everywhere. Exclusion of brent geese enhanced the level of utilisation by hares in both Festuca and Puccinellia dominated marshes, which are among the most intensively grazed parts of the salt marsh. The increase in hare grazing pressure following goose exclusion was stronger, when the adjacent control plots had attracted more goose visitation. When geese were excluded, the decrease in Festuca consumption by geese was completely matched by increased hare grazing, while for Puccinellia only part of the `surplus' was harvested. Enhanced levels of hare utilisation were not due to geese interfering directly with hare, nor due to hares avoiding goose droppings. Considering the interaction from the other perspective, hares were observed to disturb geese effectively in every spring. This might have reduced exploitation by geese of the shared resources. On the basis of our experimental results, we conclude that in this salt- marsh system competition for food with brent geese plays a role in the habitat use of hares, and that hares can reduce goose exploitation of shared habitats. Received: 30 March 1998 / Accepted: 6 July 1998     

I. Introduction

Background: Past research and ecological theory supports the hypothesis that alpine plants should be predominantly clonal, long-lived and reproduce by seed infrequently.

Aims: To challenge the entrenched view that alpine seedling establishment is uncommon and perhaps unsuccessful in maintaining alpine plant populations.

Methods: We looked for patterns in seedling establishment across a stress gradient of alpine sites and tested the notion that natural seedling recruitment would be reduced at higher altitudes due to a combination of stressful environmental conditions such as frosts, wind and extended snow cover.

Results: Seedlings were common at all study sites and there was a partial trend in mean seedling density with altitude. Seedling emergence was best predicted by a combination of altitude, plant litter and soil wilting point. Many seedlings survived to become established plants (37–61%) but there was no significant relationship between establishment and altitude. There was low similarity between the seedling flora and the standing vegetation, suggesting that patterns in seedling establishment are unlikely to be driving the patterns in the standing vegetation assemblage.

Conclusions: Although Australian alpine species predominantly reproduce vegetatively, this study shows that many plants depend on recruitment from seed in order to become established.     

Effects of Precommercial Thinning on Snowshoe Hares in Maine

Abstract: Snowshoe hares (Lepus americanus) are an important prey species and a dominant herbivore across much of their North American range, and researchers have questioned the influences of forestry practices that alter habitat for hares and the potential community-level effects on carnivores. We examined the effects of precommercial thinning (PCT) from 1 to 11 years posttreatment on snowshoe hares. In the commercial forests of northern Maine, USA, we counted and cleared hare pellets twice a year during 2001 and 2002 on >46 km of pellet transects across 30 regenerating conifer stands (17 treated with PCT) previously treated with an aerial application of herbicide. We compared densities of snowshoe hare pellets among 3 development classes with (1 yr after thinning, 6 yr after thinning, and 11 yr after thinning) and without thinning (stands with a similar history of clearcut and herbicide treatment but no thinning). During both years, densities of hares were lower in stands treated with PCT than in similar unthinned stands across the 3 development classes and during both leaf-off and leaf-on seasons (P < 0.001). Within both thinned and unthinned stands, hare density was greatest in stands in the 1-year development class when compared to the 6-year and 11-year development classes, but a statistical difference (P = 0.048) among classes was evident only during leaf-off seasons. Precommercial thinning was associated with densities of snowshoe hares that were approximately half of those in similar unthinned stands up to at least 11 years posttreatment; however, thinned stands may retain densities of hares greater than stands managed using other forest harvesting regimes. Our results apply to core portions of stands with crop trees spaced at 1.8–2.4-m intervals following complete overstory removal and herbicide treatment. We advocate caution when applying our results to other thinning regimes or across broader spatial scales.     

Uncoupled changes in tree cover and field layer vegetation at two Pyrenean treeline ecotones over 11 years

Background: The alpine treeline ecotone is regarded as a sensor of the effects of global change on alpine plant communities. However, little is known about how treeline dynamics influence the diversity and composition of alpine plant communities. Such information is necessary to forecast how ascending montane forests may affect the composition of alpine flora.

Aims: We analysed the temporal variations in tree cover, plant diversity and composition, and the effect of tree cover dynamics on field layer vegetation over a period of 11 years, at two alpine treeline ecotones in the central Pyrenees, Spain.

Methods: Tree and field layer vegetation was sampled in permanent transects in 1998 and 2009, using the point-intercept method. Temporal changes in tree cover, plant species richness and abundance were characterised along the ecotone by using a randomisation approach, rarefaction curves, and a non-parametric multivariate test, respectively.

Results: Tree cover increased significantly at one of the sites, whereas plant species richness only increased at the other site where tree cover had not changed. Vegetation composition changed significantly at both sites, but it was not spatially coupled with changes in tree cover along the ecotone.

Conclusions: A change of tree cover does not necessarily trigger changes in the ground flora at the treeline over relatively short periods (decade scale). The results challenge our ability to infer short-term biodiversity impacts from upslope advance of forests. Integrated tree and field layer monitoring approaches are necessary to produce a better understanding of the impact of ongoing global change on treeline ecotones.     

The effects of micro-habitats and grazing intensity on the vegetation of burial mounds in the Kazakh steppes

Background: Burial mounds (kurgans) of Eurasian steppes are man-made habitat islands that have the potential to harbour rich plant diversity due to micro-habitats associated with their topography.

Aims: We assessed whether kurgan micro-habitats harboured different species pools and functional groups from those found on the surrounding steppes. In addition, we asked if these mounds were affected by different grazing intensities from those on the surrounding vegetation.

Methods: We surveyed kurgan micro-habitats (northern and southern slopes, surrounding ditch) and adjacent steppe plains in non-grazed, moderately grazed and heavily grazed sites in northern Kazakhstan. We analysed differences in species composition of four habitats under three grazing regimes using Generalised Linear Mixed Models, PCA ordination and indicator species analysis.

Results: Kurgan micro-habitats had diverse vegetation and supported the co-existence of plant species with different environmental needs. We identified 16 steppe specialists confined to kurgan micro-habitats. Steppe vegetation was well-adapted to extensive grazing, although heavy grazing supported ruderals and a decline in steppe specialists. There was a significant interaction between grazing intensity and habitat type: heavy grazing supported ruderals and suppressed steppe specialists especially on the slopes.

Conclusions: We highlighted that kurgans play an important role as maintaining high plant diversity locally in extensive steppe plains in Central-Asia by increasing environmental heterogeneity and supporting specialist species confined to these micro-habitats.     

Winter Prey Selection of Canada Lynx in Northwestern Montana

ABSTRACT The roles that diet and prey abundance play in habitat selection of Canada lynx (Lynx canadensis) in the contiguous United States is poorly understood. From 1998–2002, we back-tracked radiocollared lynx (6 F, 9 M) for a distance of 582 km and we located 86 kills in northwestern Montana, USA. Lynx preyed on 7 species that included blue grouse (Dendragapus obscurus), spruce grouse (Canachites canadensis), northern flying squirrel (Glaucomys sabrinus), red squirrel (Tamiasciurus hudsonicus), snowshoe hare (Lepus americanus), least weasel (Mustela nivalis), and white-tailed deer (Odocoileus virginianus). Snowshoe hares (69 kills) accounted for 96% (4-yr average, range = 94–99%) of prey biomass during the sample period. Red squirrels were the second-most-common prey (11 kills), but they only provided 2% biomass of the winter diet. Red squirrels contributed little to the lynx diet despite low hare densities. A logistic regression model of snowshoe hare, red squirrel, and grouse abundance, as indexed by the number of track crossings of use and available lynx back-tracks, was a significant (Wald statistic = 19.03, df = 3, P < 0.001) predictor of habitat use. As we expected, lynx (P < 0.001) selected use-areas with higher snowshoe hare abundance compared to random expectation. However, the red squirrel index had a weak (P = 0.087) negative relationship to lynx use, and grouse was nonsignificant (P = 0.432). Our results indicate that lynx in western Montana prey almost exclusively on snowshoe hares during the winter with little use of alternative prey. Thus, reductions in horizontal cover for hares would degrade lynx habitat.     

Positive effects of an extremely hot summer on propagule rain in upper alpine to subnival habitats of the Central Eastern Alps

Background: Propagule production and dispersal largely determine the distribution and potential migration ability of alpine plant species. Variation in reproductive success caused by year-to-year variation in climate may critically influence these processes.

Aims: To obtain estimates for the propagule rain in high-alpine plant communities and detect potential dispersal events from lower elevations.

Methods: The magnitude and composition of the propagule rain was studied in different plant communities along an elevation gradient from the upper alpine to subnival zone. Propagules were trapped at eight elevations from 2760 to 3070 m a.s.l. for three years from July to September 2003–2005. Vascular plant species and their cover were recorded in an area with a radius of 10 m surrounding the traps.

Results: A five- to 10-fold higher propagule rain was observed in 2003, a year with an exceptionally hot summer, compared to 2004 and 2005. Propagule and species numbers varied highly among years and community types. Few propagules of non-local origin were recorded in any year.

Conclusions: Extremely hot summers are likely to greatly magnify the propagule rain size of species in alpine habitats. Such ‘mast years’ may contribute to enhanced and accelerated vegetation changes in alpine habitats in the absence of limiting factors.     

Is Size of Fecal Pellets a Reliable Indicator of Species of Leporids in the Southern Rocky Mountains?

Abstract: In the southern Rocky Mountains, USA, snowshoe hares (Lepus americanus) and mountain cottontails (Sylvilagus nuttallii) are syntopic. Previous researchers used size of fecal pellets to identify leporid species in the Southern Rockies, a common criterion for identifying species of cervids. We measured 655 fecal pellets from 10 mountain cottontails and 2,374 fecal pellets from 23 snowshoe hares. We found no relationship between the body weight of mountain cottontails and the size of their fecal pellets (r = 0.04, F = 0.01, P = 0.91) but found one for snowshoe hares (r = 0.48, F = 9.3, P = 0.005). Although the 2 species differed in the size of their fecal pellets, the difference between means (1.2 mm) was sufficiently small to require measuring individual pellets and is only applicable to individuals of adult size. Although fecal pellet counts may be used to estimate presence and relative abundance of snowshoe hares in the absence of syntopic leporids, where multiple species of leporids are syntopic this method may yield misleading results.     

Winter Habitat Selection by Canada Lynx in Maine: Prey Abundance or Accessibility?

Abstract: We related winter habitat selection by Canada lynx (Lynx canadensis), relative abundance of snowshoe hares (Lepus americanus), and understory stem densities to evaluate whether lynx select stands with the greatest snowshoe hare densities or the greatest prey accessibility. Lynx (3 F, 3 M) selected tall (4.4-7.3 m) regenerating clear-cuts (11-26 yr postharvest) and established partially harvested stands (11-21 yr postharvest) and selected against short (3.4-4.3 m) regenerating clear-cuts, recent partially harvested stands (1-10 yr), mature second-growth stands (>40 yr), and roads and their edges (30 m on either side of roads). Lynx selected stands that provided intermediate to high hare density and intermediate cover for hares (i.e., prey access) but exhibited lower relative preference for stand types with highest hare densities where coniferous saplings exceeded 14,000 stems/ha.     

A review of the status and decline in abundance of the Irish Hare (Lepus timidus hibernicus) in Northern Ireland

The Northern Ireland Hare Survey documented the distribution of the Irish Hare (Lepus timidus hibernicus). Historical game bag records and other, more contemporary, records of hare distribution were examined. These data indicate how numbers of L. t. hibernicus may have changed over the last 140 years. The results of the Northern Ireland Hare Survey suggested that L. t. hibernicus was widespread throughout Northern Ireland. Current average densities are no more than 0.65 hares/km². Game bag records indicate that hare densities may have been much higher in the past, with a maximum of 138 hares/km² recorded on Crom Estate, Co. Fermanagh, in 1864. Evidence from hare distribution recorded during the Northern Ireland Rabbit Survey indicates that hare numbers declined between 1984 and 1994. Evidence from all sources suggests that L. t. hibernicus has declined in abundance substantially, with present total population estimates for Northern Ireland ranging from 8250 to 21 000 individuals. Flushing data indicate that rushes and hedgerows are important diurnal resting areas for hares. While the principal reason for the decline in numbers of L. t. hibernicus in Northern Ireland is not clear, more species‐rich pasture and provision of areas of cover, such as rushes, may arrest further declines, or indeed promote numbers of hares, particularly in lowland areas.     

Influence of forest structure on the abundance of snowshoe hares in western Wyoming

Snowshoe hares (Lepus americanus) are a primary prey species for Canada lynx (Lynx canadensis) in western North America. Lynx management plans require knowledge of potential prey distribution and abundance in the western United States. Whether even-aged regenerating forests or multi-storied forests contain more snowshoe hares is currently unknown. During 2006–2008, we estimated snowshoe hare density in 3 classes of 30–70-year-old lodgepole pine (Pinus contorta) and 4 classes of late seral multi-storied forest with a spruce (Picea engelmannii)-fir (Abies lasiocarpa) component in the Bridger-Teton National Forest, Wyoming. We recorded physiographic variables and forest structure characteristics to understand how these factors influence abundance of snowshoe hares. In many instances, snowshoe hares were more abundant in late seral multi-storied forests than regenerating even-aged forests. Forest attributes predicting hare abundance were often more prevalent in multi-storied forests. Late seral multi-storied forests with a spruce–fir component and dense horizontal cover, as well as 30–70-year-old lodgepole pine with high stem density, were disproportionately influential in explaining snowshoe hare densities in western Wyoming. In order to promote improved habitat conditions for snowshoe hares in this region, management agencies should consider shifting their focus towards maintaining, enhancing, and promoting multi-storied forests with dense horizontal cover, as well as developing 30–70-year-old lodgepole pine stands with high stem density that structurally mimic multi-storied forests. © 2012 The Wildlife Society.     

The Reaction of the Medium in Relation to Root Formation in Coleus

Background: As seed dispersal can vary among years and individuals, studies that focus on a single year or on a few individuals may lead to erroneous conclusions.

Aims: To study temporal and spatial intraspecific variation of seed dispersal in Scrophularia canina, a widespread species with capsule-type fruit.

Methods: Primary seed dispersal was quantified by placing traps in each cardinal direction around 10 individuals during two consecutive years. We correlated several seed shadow parameters (modal dispersal distance, kurtosis, skewness, percentiles, slope, and seed percentage beneath the plant canopy) with three plant features (maximum height, lateral spread and seed production).

Results: Scrophularia canina dispersed their seeds by boleochory, giving rise to a typical leptokurtic curve, but behaving as a barochorous species, because about 90% of seeds landed beneath the plant canopy. Temporal dispersal in S. canina included several seed waves associated with maximum wind speeds. Plant lateral spread was significantly positively correlated with seed percentiles and percentage of seeds beneath the plant canopy regardless of year. A seed production effect was only evident when both years were considered together.

Conclusions: Although time-consuming, investigation of the dispersal process for more than 1 year provides more realistic information on seed dispersal. Lateral spread is the main plant feature determining seed shadow.