Lignin signatures show the effects of changes in heather and bracken cover on the composition of organic matter in a moorland soil profile

Bracken (Pteridium aquilinum) is aggressively displacing heather (Calluna vulgaris) on many moorlands in Britain. We investigated the use of lignin derivatives to identify the distribution of soil organic matter (SOM) derived from bracken in moorland soil profiles formed under heather. Phenylpropanoids extracted from recently senesced litters, roots and SOM, using alkaline CuO oxidation, showed distinct signatures for bracken and heather, with vanillyl moieties dominating bracken litter extracts and vanillyl and syringyl dominating heather litter extracts. Ratios of vanillyl and syringyl concentrations characterised the SOM derived from heather and bracken better than the concentrations of the individual moieties. The analysis showed up to a depth of 5 cm under pure bracken cover, and at the interface between heather and bracken, the SOM was largely derived from bracken litter but below that depth SOM was apparently derived from heather. The use of these methods to identify the plant origin of SOM not only enables understanding the effects of changing vegetation cover on organic matter dynamics in moorland soils but could also facilitate management techniques in moorland/heathland restoration which involve the removal of comparatively nutrient-rich SOM derived from bracken. Received: 30 November 1997 / Accepted: 11 April 1998     

The morphology of bracken (Pteridium aquilinum (L.) Kuhn) in the North York Moors—a comparison of the mature stand and the interface with heather (Calluna vulgaris (L.) Hull) 1. The fronds

The encroachment of bracken (Pteridium aquilinum (L.) Kuhn) into areas previously dominated by heather represents a threat to the ecology, agricultural economy and landscape value of many UK upland areas, including the moorland of the North York Moors National Park. The morphology of bracken, within a mature stand and at several bracken-heather interfaces, has been studied at a number of sites within the National Park. Differences have been found in the frond growth of bracken in a mature stand, at stationary stand margins, and at advancing stand margins where bracken is encroaching into heather. Frequency of fronds present on bracken rhizome growing at a stationary stand margin close to the interface with heather (1–2 m behind the boundary) are approximately the same as those found within a mature stand. At advancing margins (again 1–2 m behind the boundary), maximum frond densities were often found to exceed those present in either a mature stand or at a stationary margin. Frond numbers decline rapidly at the stand margins as distance from the stand increases. This is especially true where the front is stationary and bracken is not encroaching into heather at a significant rate. Maximum frond heights in a mature stand consistently exceed those at stand margins (even 1–2 m into the stand) and are greater at stationary margins than at advancing margins. Outlying fronds at the edges of bracken stands are generally present in greater numbers, and further into the area dominated by heather, where the margin is advancing. Heights of outlying fronds fall as distance from the bracken stand increases, as does stipe length. Fronds at the edges of bracken stands emerge each spring before those further into the stand and are therefore particularly vulnerable to frost damage. Outlying fronds are not, however, the first to emerge. Early emerging fronds reach their maximum height and eventually become senescent before later emerging fronds. Whilst most fronds emerge before the end of June a few fronds continue to emerge throughout the summer. Frond densities close to the edges of bracken stands (1–2 m into the stand) are comparable to those in a mature stand. At advancing stand margins frond densities generally exceed those in a mature stand, suggesting that a large number of potential entry points for foliage-applied herbicides are available for bracken control at the stand margins. The ratio of potential uptake points to biomass of rhizome is also greatest at the edges of the stand, and the canopy 1–2 m into the stand is usually almost completely closed. It is possible therefore, the efficacy of herbicides could be improved by the use of small scale applications, using tractors or hand-held sprayers, close to the margins of bracken stands.     

Responses of bracken to increased temperature and nitrogen availability

In 1993 a project was set up at Upper Teesdale to investigate some of the effects of predicted increased temperatures and increased nutrient availability resulting from increased litter decomposition rates, and atmospheric deposition of nitrogen, on the interaction between bracken and heather, two species which cover large areas globally. 32 2 × 2 m plots were laid out on a hillside on, and just below, the interface between the two species. Half of the plots were placed on pure bracken, and the other half on the boundary. Within each vegetation type half of the plots were covered with open-topped polythene tents to simulate climate warming, while the other plots remained open. The second treatment consisted of additional fertilizer at 50 kg N ha⁻¹ y⁻¹ simulating increased decomposition rates at higher temperatures and increased atmospheric nitrogen deposition. The two treatments and their controls were combined in a 2 × 2 factorial design, with two replicates in each of two blocks. Results obtained in the first two summers indicated that fronds emerged earlier in spring and senesced later in autumn inside the tents, effectively lengthening the growing season. Fronds were taller when growing in warmer conditions, and carried more pinnae. In addition, the frond density was higher inside the tents, and together these effects contributed to greatly enhanced vigour of bracken in some of the conditions predicted by General Circulation Models. It seems likely that bracken will be more competitive, and have the potential to further encroach into heather dominated areas under these conditions.     

The morphology of bracken (Pteridium aquilinum (L.) Kuhn) in the North York Moors—a comparison of the mature stand and the interface with heather (Calluna vulgaris (L.) Hull) 2. The rhizome

The rhizome system of mature bracken (Pteridium aquilinum (L.) Kuhn) contains large reserves of both biomass (mean 8.63 kg m?² fr. wt) and buds (mean 565 m^-2) which are largely responsible for both its persistence and its often rapid rates of vegetative encroachment. Within areas such as the North York Moors the spread of bracken into areas previously dominated by heather and grass is considered undesirable because of reduced land value in terms of both agriculture and ecological diversity. In this paper we describe the morphology of bracken rhizome within a mature bracken stand, and at advancing and stationary stand margins where bracken-heather interfaces occur. Stationary margins, i.e. those where bracken is not encroaching into heather at a significant rate, often have morphological characteristics intermediate to those of a mature stand and an advancing margin. In the mature stand rhizome biomass is dominated by carbohydrate-storing long shoots which comprise 63% of the total fresh weight, whilst the majority of rhizome buds (89% of all active and 86% of all dormant buds) are found on frond-bearing short shoots. At the margins of a bracken stand the proportion of rhizome which is composed of long shoots is even greater, and that of short shoots small relative to that in the mature stand. More transitional shoots are also found at the stand margins. Hence close to the margin a greater proportion of fronds is found on transitional rhizome than is the case in more mature parts of the stand. The majority of buds on all types of rhizome are in a dormant state. The proportion of buds which are active is, however, greater on long and transitional shoots than on short shoots. Hence, a larger proportion of buds are active close to the margin where the rhizome is composed less of short shoots than is the case further into a mature stand. The differences in the morphology of bracken in a mature stand and at the stand margins which are identified here support the idea of controlling bracken at stand margins in preference to the spraying of large areas of dense, mature bracken. Morphological differences include an increased proportion of active buds, greater frequency of fronds per unit rhizome biomass, reduced biomass reserves. Improved conditions for the re-invasion and re-establishment of alternative vegetation are also available at stand margins in comparison with the centre of a dense bracken stand.     

Bracken-induced increase in soil P availability,along with its high P acquisition efficiency,enables it to invade P-deficient meadows

欧洲蕨入侵提高了土壤磷有效性和高磷获取效率 对于欧洲蕨(Pteridium aquilinum)入侵荒地后土壤化学的变化,已经有了相关的研究,但是对于欧洲蕨入侵草地的研究却比较缺乏。本研究探讨了欧洲蕨入侵缺磷草地是否会改变土壤养分资源库,以及与土壤过程和欧洲蕨营养相关的机制。此外,还研究了欧洲蕨入侵前后群落组成对土壤化学差异的反应。在蕨菜生物量高峰期间,我们进行了土壤和植物取样以及植被调查。数据分析包括方差分析(ANOVA)和典型对应分析(CCA)。研究结果表明,欧洲蕨入侵提高了土壤磷有效性、土壤有机碳浓度以及碳氮比、碳硫比和氮硫比,同时降低了铁和钴的浓度。欧洲蕨羽片富含磷,根茎富含钾,而羽片 和根茎的氮磷比很低。典型对应分析(CCA)显示了与磷和钾有效性相关的常见草地植物物种的不同丰度模式。在磷有效性极低的条件下,绒毛草(Holcus lanatus)表现出竞争优势。欧洲蕨通过促进铁和铝的浸出提高了磷有效性。从土壤资源生态位的角度来看,欧洲蕨通过提高自身生长所需的磷有效性并且增加对其他物种的氮限制,从而获得了竞争优势。欧洲蕨提高土壤磷有效性的能力,以及其高磷获取效率背后的生理机制,似乎将欧洲蕨与其他竞争生态策略的物种区分开来,因为其他物种的生长主要局限于营养丰富的环境,因此欧洲蕨更容易入侵缺磷的草地。     

The Use of Mixtures of Sulfur and Bracken Litter to Reduce pH of Former Arable Soils and Control Ruderal Species

High soil pH has been highlighted as a constraint to the restoration of heathland on ex‐arable land. Previous studies at the Minsmere Reserve of the Royal Society for the Protection of Birds (RSPB) in England have shown that it is possible to acidify ex‐arable soils using elemental sulfur and bracken litter, although sulfur (S) is more effective. Current recommendations suggest that 4 tS/ha need to be applied to reduce soil pH below pH 4, control vigorous ruderal species, and create conditions suitable for Calluna vulgaris (heather) establishment. However, S is relatively expensive, and as bracken litter is moderately abundant within the reserve, it made economic sense to evaluate the potential for mixing S with bracken to see if adequate pH reductions could be achieved at lower S rates. Accordingly an experiment was designed to test the effects of combining S (0–8 t S/ha) and bracken litter (0–10 cm depth layers) on (1) soil pH, (2) cover of ruderal species, and (3) the developing plant community. Significant interactions were detected, especially in the period immediately after application. Where bracken litter was applied the soil pH fell immediately; in contrast, S took at least six months to start reducing pH. Where mixtures were applied there was a synergistic effect, which produced a lower pH than the S or bracken litter applied alone. These effects were most marked at low S application rates, between 0.5–4 t S/ha. The effects of the bracken litter addition also reduced the growth of ruderal species in the period immediately after application, probably through a combination of acidification and physical smothering. There is, therefore, a clear potential to acidify ex‐arable soils using combinations of S and bracken litter in schemes designed to restore Calluna heathland.     

Effects of nutrient supply,light availability and herbivory on the growth of heather and three competing grass species

Grasses are becoming more abundant in areas in NE Scotland which until recently were dominated by heather (Calluna vulgaris). However, it is not clear if grasses are aggressive competitors which are now able to outcompete the dwarf shrub due to changes in environmental factors (such as grazing pressure and increasing nutrient inputs), or just opportunistic invaders, occupying gaps in the canopy which occur when heather reaches the degenerate stage. Experiments in turves and in field plots were carried out in order to investigate the performance of three grass species, Nardus stricta, Deschampsia cespitosa and Deschampsia flexuosa growing in competition with heather. These three species were selected because they differ in their nutrient requirements, palatability to herbivores and tolerance of shading. The grasses were planted in heather canopies of different structure, either turves of heather of different height and age, or moorland plots with or without heavy grazing by sheep and deer. Fertiliser (NPK) was applied to half the experimental plants. The growth of the grass species and the heather in response to the fertiliser and grazing treatments was measured, together with the light levels penetrating the canopy and received by the grass plants.Results indicated that heather was likely to be outcompeted by grasses only when there are gaps in the canopy, resulting either from heavy grazing or from the heather being in the mature or degenerate phase. Fertiliser enhanced plant growth whereas fencing out herbivores led to strong competition for light as the heather canopy closed. It is concluded that grasses require gaps in the canopy to successfully invade heather moorland, or they tend to be shaded out. Thus better management of heather moorlands to maintain a dense canopy structure may help to preserve heather cover even under increasing nutrient inputs.     

Utilisation of different ages of heather on three Scottish moors by red grouse, mountain hares, sheep and red deer

Utilisation of heather at three Scottish moors, by red grouse, mountain hares, sheep and red deer, was measured by collecting their faecal droppings regularly from plots on heather patches of different ages. Rates of heather growth differed between moors, and preferences of the four species, all grazers of heather, were related more closely to its height than to its age. Thus hares spent most time on ground with heather less than 15 cm high, sheep less than 20 cm, grouse preferred heather 10–30 cm high and deer spent most time on heather over 25 cm high.
Differences between outer, middle and inner plots on patches at one of the moors in dicated that grouse are reluctant to move far from cover into areas of newly burned ground, and reluctant to move far into patches of tall, dense heather, when these are next to shorter material. Hares preferred the middle of patches on newly burned ground and very young heather.
At two of the moors, but not the third, there was evidence that heavy grazing by hares in winter reduced the amount of young heather available to hares and grouse in spring. This potential reduction in the quality of the spring diet of grouse is discussed In relation to previously reported associations between numbers of hares and grouse at those two moors.     

Effects of environmental change, including drought, on water use by competing Calluna vulgaris (heather) and Pteridium aquilinum (bracken)

1. Competition for water between Calluna vulgaris (heather) and Pteridium aquilinum (bracken) was studied in conditions of increased temperature, drought and increased nitrogen supply. All these factors increased the intensity of competition for water, with the combination of drought and increased nitrogen having the greatest effect on water use.
2. Both species increased water-use efficiency in response to increased nitrogen and drought. The effects of temperature were however, equivocal. Calluna had a greater water demand than Pteridium but acclimated to water stress more readily.
3. Calluna was the superior competitor for water; its water-use efficiency was reduced as a consequence of its roots depleting water from the Pteridium rooting zone. Pteridium , the poorer competitor, increased water-use efficiency to cope with reduced water availability owing to competition.
4. There was a strong relationship between carbon isotope discrimination (Δ) and instantaneous water-use efficiency for both species, but discrimination provided a more sensitive measure of seasonal water-use efficiency. Reconstruction of the plant's history of water-use efficiency by retrospective measurement of Δ proved a useful technique for Calluna leaves but was inappropriate for Pteridium rhizome.     

Densities of Ixodes ricinus ticks (Acari: Ixodidae) on moorland vegetation communities in the UK

Unfed (questing) Ixodes ricinus ticks were collected by blanket dragging on a monthly basis from heather-dominated, Vaccinium-dominated and bracken-dominated vegetation communities from two different biogeographical regions of the UK (the Quantock Hills in Somerset, south west England and the North York Moors, north east England) throughout the spring and summer months of 1991 and 1992. Eighteen sites were monitored across the two regions and a total of 1920 blanket drags were carried out. Vaccinium sites showed high tick densities at all life stages, as did bracken sites. Significantly lower numbers of larval and nymphal ticks per drag were collected on heather sites than were collected on either Vaccinium (bilberry/whortleberry) or bracken sites, while similar numbers of adult ticks per drag were collected from each of the three vegetation communities. There was no significant difference between the mean numbers of any tick life stage collected on the Quantock Hills and those collected on the North York Moors on these vegetation communities or between the mean numbers of any tick life stage collected in 1991 and those collected in 1992 on these vegetation communities.     

Bracken invasion in Scotland

Summary

For the first time, estimates of the amount of sparse bracken on open land, the amounts of bracken under tree cover and the amounts in linear features are made. These data were used to predict the areas of Scotland most at risk from invasion from patch edges, from sparse patches becoming dense and from invasion along linear features. Computer models show that if the climate warms over the next 50 yr as predicted, then bracken growth will increase and the amount of hill land suitable for invasion by bracken will increase. On some sites where bracken is being controlled, it is reduced in cover and other ground vegetation is becoming established. However, we cannot ensure that when bracken is controlled that the control is always effective and that appropriate vegetation will become established. Some generalized rules have been derived to help in the selection of sites likely to show the most benefit from control treatment.     

Simulation of bracken cover in forested areas in Victoria in response to season,overstorey and fire conditions

Abstract A computer-based model is presented that brings together the existing knowledge about the dynamics of bracken in eucalypt forests in south-eastern Australia. The average length of the growing season, the timing and extent of logging, and the timing and severity of fire are major inputs to the model. The model is predominantly deterministic, but has a stochastic environmental component. The model is validated with four independent sets of data. The model successfully predicts the trends in the data. The interactions between bracken, other understorey species and eucalypt regeneration are explored with the model. The likely effect of different silvicultural and fire management practices on the cover of bracken are investigated using the model. Year to year variation can mask the effects of disturbance if bracken cover is viewed in any one year in isolation. Bracken cover trends can only be seen over a period of 10 years or more. The model reinforces the importance of site productivity as measured by length of growing season and the degree of tree removal on bracken cover.     

The history and ethnobotany of bracken

The fossil record shows that Pteridium has been present in all the temperate stages of the Quaternary, but that for most of this period it formed a component of the herb layer of deciduous woodlands. Bracken did not begin to reach its present abundance and importance until widespread forest clearance began with the arrival of Neolithic man about 5000 years ago. Some of the factors that have played a role in the history of bracken are illustrated by a discussion of the spread of bracken in Scotland that occurred contemporaneously with the change from cattle- to sheep-farming in the 18th century. It is shown that man has played a dominant role in influencing the spread of bracken in Scotland, but that bracken was considered of great positive economic value. The ethnobotany of bracken is discussed, and it is suggested that it was once an important source of potash for the glass, soap and bleaching industries. The use of the plant as fuel, thatch, litter, compost and food, and for medicinal purposes is considered, and mention is made of various minor uses of the plant. Unfortunately it is difficult to estimate the amount of bracken consumed by these various employments, but it is suggested that the increased abundance of the plant may have been exaggerated because what was once a useful resource has now become a pest.     

The ecological implications of bracken control to plant/animal systems

The importance of considering bracken as part of a dynamic soil/plant/animal system is emphasized and the extent, distribution and successional relationships of the fern in the uplands are discussed. Pteridietum is classified into six types according to its pattern of local distribution. Bracken communities as animal habitat are considered in relation to distribution types, frond density and canopy cover. As a direct source of food for the vertebrates bracken is considered to be of little or no value. As an indirect source of food bracken communities may be important in providing habitats for invertebrates, though in terms of numbers of species the insect fauna of vigorous Pteridietum is not great compared with that of other communities that may replace it. Pteridietum provides shelter and cover for a number of small mammals but in these respects it is of very limited value to the larger grazing animals. It is estimated that in Scotland, under the economic conditions of recent years, bracken herbicide is unlikely to be used on more than about 40,470 ha (100,000 acres) in the combined land uses of agriculture, forestry and sport; i.e. less than 25% of the Scottish bracken coverage is at risk. At this level of control, provided some precautions are taken, it is considered that the wild vertebrates are unlikely to be seriously affected; indeed in certain situations some benefits may be derived.     

Model parameterization to simulate and compare the PAR absorption potential of two competing plant species

Mountain pastures dominated by the pasture grass Setaria sphacelata in the Andes of southern Ecuador are heavily infested by southern bracken (Pteridium arachnoideum), a major problem for pasture management. Field observations suggest that bracken might outcompete the grass due to its competitive strength with regard to the absorption of photosynthetically active radiation (PAR). To understand the PAR absorption potential of both species, the aims of the current paper are to (1) parameterize a radiation scheme of a two-big-leaf model by deriving structural (LAI, leaf angle parameter) and optical (leaf albedo, transmittance) plant traits for average individuals from field surveys, (2) to initialize the properly parameterized radiation scheme with realistic global irradiation conditions of the Rio San Francisco Valley in the Andes of southern Ecuador, and (3) to compare the PAR absorption capabilities of both species under typical local weather conditions. Field data show that bracken reveals a slightly higher average leaf area index (LAI) and more horizontally oriented leaves in comparison to Setaria. Spectrometer measurements reveal that bracken and Setaria are characterized by a similar average leaf absorptance. Simulations with the average diurnal course of incoming solar radiation (1998–2005) and the mean leaf–sun geometry reveal that PAR absorption is fairly equal for both species. However, the comparison of typical clear and overcast days show that two parameters, (1) the relation of incoming diffuse and direct irradiance, and (2) the leaf–sun geometry play a major role for PAR absorption in the two-big-leaf approach: Under cloudy sky conditions (mainly diffuse irradiance), PAR absorption is slightly higher for Setaria while under clear sky conditions (mainly direct irradiance), the average bracken individual is characterized by a higher PAR absorption potential. (∼74 MJ m⁻² year⁻¹). The latter situation which occurs if the maximum daily irradiance exceeds 615 W m⁻² is mainly due to the nearly orthogonal incidence of the direct solar beam onto the horizontally oriented frond area which implies a high amount of direct PAR absorption during the noon maximum of direct irradiance. Such situations of solar irradiance favoring a higher PAR absorptance of bracken occur in ∼36% of the observation period (1998–2005). By considering the annual course of PAR irradiance in the San Francisco Valley, the clear advantage of bracken on clear days (36% of all days) is completely compensated by the slight but more frequent advantage of Setaria under overcast conditions (64% of all days). This means that neither bracken nor Setaria show a distinct advantage in PAR absorption capability under the current climatic conditions of the study area.     

The effect of glyphosate on frond regeneration, bud development and survival, and storage rhizome starch content in bracken

In addition to very effectively suppressing bracken frond regeneration for two years at levels of 2.24 kg a. i./ha and above, glyphosate had considerable effects on the rhizome system. Dry weight of frond-bearing rhizomes, number of living apices and developed buds, number and viability of dormant buds and starch content of storage rhizomes were all markedly reduced. The effects on dormant buds and starch content are particularly important when considering frond regeneration and as a consequence of these, glyphosate is likely to give long lasting control of bracken. Low levels of glyphosate (0.02 and 0.07 kg a. i./ha) did not stimulate bracken growth, while the addition of NH₄SCN apparently inhibited the action of the sub-lethal rate of 0.56 kg a. i./ha.     

Land‐use Change Dynamics,Soil Type and Species Forming Mono‐dominant Patches: the Case of Pteridium aquilinum in a Neotropical Rain Forest Region

Deforestation and agricultural land degradation in tropical regions can create conditions for growth of perennial plant species forming mono‐dominated patches (MDP). Such species might limit forest regeneration, and their proliferation forces the abandonment of fields and subsequent deforestation to establish new fields. Therefore, identifying factors fostering MDP species is critical for biodiversity conservation in human‐modified landscapes. Here, we propose a conceptual framework to identify such factors and apply it to the case of Pteridium aquilinum (bracken fern), a light‐demanding species, tolerant of low soil fertility and fire. We hypothesize that bracken proliferation is promoted by land‐use changes that increase light availability, especially in sites with low soil fertility and land uses involving fire. We assessed this idea using agricultural fields in southeastern Mexico with different land‐use change histories and quantifying prevalence and cover of bracken. Five different land‐use change histories resulted from transitions among forest, crop, pasture, and fallow field stages. Of the 133 fields sampled, 71 percent had P. aquilinum; regression tree analysis indicated that 65 percent of inter‐field variation in prevalence and 90 percent in cover was explained by land‐use change history and soil type. Maximum prevalence, cover, and rates of increase in bracken were found on fields with low fertility sandy/clay soils, which had been used for crops and pasture, were frequently burned, and had high levels of light. Fields on fertile alluvial soil never used for pasture were bracken‐free. Agriculture promoting high light environments on less fertile soils is a major cause of bracken proliferation and likely that of other MDP species.     

Bracken, ants and extrafloral nectaries. III. How insect herbivores avoid ant predation

Abstract. 1. Ants (Myrmica spp. and Formica lemani) visiting the extrafloral nectaries of bracken, Pteridium aquilinum, imposed heavy mortality on caterpillars of a novel moth species experimentally introduced onto bracken fronds. However, the exclusion of ants from bracken fronds had no significant effect on adapted bracken-feeding herbivores in Britain.
2. The feeding stages of British bracken-feeding insects are immune to, or can largely avoid, predation by ants in a variety of ways. Gall formers and miners cannot be attacked by these ants. Two other species hide, one inside tied leaves, the other in a mass of 'spittle'. Another group of species jumps away from, or falls off the plant when touched by ants. Sawfly caterpillars in the genera Strongylogaster, Aneugmenus and Tenthredo have viscous, distasteful haemolymph that repels ants.
3. No species of bracken herbivore has an absolute temporal refuge from ants; during their development they all overlap with ants to some degree.
4. Ant predation appears to have played a significant role in determining the contemporary structure of British bracken-feeding insect communities. Distasteful haemolymph in sawfly caterpillars may have evolved in response to selection from ant predation. Other species may fortuitously possess characteristics, evolved in response to a variety of selective forces, that also reduce the impact of ants; without such characteristics, however, we postulate that they would be unable to live on this plant. An absence of external, foliage feeding Lepidoptera early in the spring, a high proportion of sawfly species, and a high proportion of gall-formers and miners may all be characteristics of the bracken herbivore community which have been influenced by ant predation.     

Strategies for nematode transmission: selective migration of Trichostrongylus tenuis infective larvae

Successful transmission of macroparasites is dependent on exposure of susceptible hosts to free-living infective stages. When these hosts are herbivores that feed mostly on a single food plant then natural selection should favour those infective larvae that selectively ascend this main food plant. Red grouse feed predominantly on heather, Calluna vulgaris, so we predict that the infective larvae (L3) of the caecal nematode Trichostrongylus tenuis selectively locate and ascend heather plants. To determine whether the presence of heather influences the horizontal dispersal of T. tenuis L3 across soil, the movement of L3 across trays of soil with and without heather was investigated in the laboratory. More T. tenuis L3 were recovered from soil when heather was present, implying that larval migration may be influenced by chemical cues produced by heather plants. This was investigated in a second experiment, in which the horizontal dispersal of T. tenuis larvae was examined in the presence of heather and grass vegetation. This trial was repeated with larvae of a second species, Haemonchus contortus, a nematode whose hosts feed on a wide range of grass and shrub species. Significantly more larvae of both nematode species were recovered in the region of the heather than the grass or controls. This implies that T. tenuis and H. contortus L3 exhibit selective migration towards heather, perhaps reflecting a general response to plant cues which may be stronger for heather than for grass.     

Tree seed rain and seed removal,but not the seed bank,impede forest recovery in bracken (Pteridium aquilinum (L.) Kuhn)‐dominated clearings in the African highlands

Considerable areas dominated by bracken Pteridium aquilinum (L.) Kuhn occur worldwide and are associated with arrested forest recovery. How forest recovery is impeded in these areas remains poorly understood, especially in the African highlands. The component processes that can lead to recruitment limitation—including low seed arrival, availability and persistence—are important determinants of plant communities and offer a potential explanation for bracken persistence. We investigated key processes that can contribute to recruitment limitation in bracken‐dominated clearings in the Bwindi Impenetrable National Park, Uganda. We examined if differences in seed rain (dispersal limitation), soil seed bank, or seed removal (seed viability and persistence) can, individually or in combination, explain the differences in tree regeneration found between bracken‐dominated areas and the neighboring forest. These processes were assessed along ten 50‐m transects crossing the forest–bracken boundary. When compared to the neighboring forest, bracken clearings had fewer seedlings (bracken 11,557 ± 5482 vs. forest 34,515 ± 6066 seedlings/ha), lower seed rain (949 ± 582 vs. 1605 ± 335 tree seeds m^?2 year^?1), comparable but sparse soil seed bank (304 ± 236 vs. 264 ± 99 viable tree seeds/m²), higher seed removal (70.1% ± 2.4% vs. 40.6% ± 2.4% over a 3‐day interval), and markedly higher rodent densities (25.7 ± 5.4 vs. 5.0 ± 1.6 rodents per 100 trapping sessions). Camera traps revealed that rodents were the dominant animals visiting the seeds in our seed removal study. Synthesis: Recruitment limitation contributes to both the slow recovery of forest in bracken‐dominated areas, and to the composition of the tree species that occur. Low seed arrival and low persistence of unburied seeds can both explain the reduced density of seedlings found in bracken versus neighboring forest. Seed removal, likely due to rodents, in particular appears sufficient to constrain forest recovery and impacts some species more severely than others.