Interaction of herbivory and seasonality on the dynamics of Caribbean macroalgae

Many Caribbean coral reefs are undergoing a phase shift from coral to macroalgal dominance. Understanding the processes driving changes in algal abundance and community structure requires clarification of the relative effects of top-down (e.g., herbivory) and bottom-up processes (e.g., light, temperature, and nutrients). To date, a number of studies have examined the relative effects of grazing versus nutrification but interactions between herbivory and natural, seasonal fluctuations in temperature and light have not been investigated. This study considered the dynamics of three Caribbean macroalgal species [Lobophora variegata (Lamouroux), Dictyota pulchella (H?rnig and Schnetter), and Halimeda opuntia (Linnaeus)] and algal turf. A field experiment was established to measure species-specific algal dynamics (changes in abundance) over 13?months in the presence and absence of herbivory. Both herbivory and seasonal changes were important processes controlling macroalgal and turf abundance. Water temperature and light had a key role on D. pulchella; this species?? abundance significantly increased in the summer, when water temperature and light were the highest, and decreased during winter. Surprisingly, herbivory did not seem to control D. pulchella directly. However, herbivory was the most important process controlling the abundance of L. variegata, H. opuntia, and turf. The abundance of both algal species was correlated with seasonal changes in the environment, but was depleted outside cages throughout the year. The abundance of H. opuntia was positively correlated with temperature and light, but there was no statistical interaction between drivers. The statistical interaction between temperature and light was significant for the abundance of L. variegata and turf, but algal abundance declined as both factors increased. Overall, macroalgal and turf cover were mainly controlled by herbivory, while community structure (which species contributed to the overall cover) was largely influenced by seasonal changes in temperature and light.     

Patch dynamics of the stream macrophyte, Callitriche cophocarpa

1. We examined changes in position and growth of Callitriche cophocarpa patches in a shallow section of a small Danish stream during the main expansion of plant cover between April and June. Mean upstream growth of patches was only 7.5% of downstream growth. The mean growth rate was 1,02cm day^?1 in length and 0.55 cm day^?1 in width and the growth rates were not significantly dependent on patch size. For patches with an area (A) above 100cm², the growth rates in patch area (dA/dt) was proportional to patch circumference and thus √Aand the relative growth rate (dA/dt A^?1) was inversely proportional to √A.The smallest patches (<100cm²), however, expanded less than expected because of their combined tendency to grow more slowly in patch length and width than larger patches. 2. The expansion of plant cover will be much more rapid in many medium-sized patches compared to few large patches of a similar combined area, because internal selflimitation by light and space is partially relieved in smaller patches. We anticipate that recruitment and mortality of new-formed patches are critical steps preceding areal expansion. Quantification of these processes are needed fully to evaluate patch dynamics and space occupation.     

Allelopathy in the tropical alga Lobophora variegata (Phaeophyceae): mechanistic basis for a phase shift on mesophotic coral reefs?

Macroalgal phase shifts on Caribbean reefs have been reported with increasing frequency, and recent reports of these changes on mesophotic coral reefs have raised questions regarding the mechanistic processes behind algal population expansions to deeper depths. The brown alga Lobophora variegata is a dominant species on many shallow and deep coral reefs of the Caribbean and Pacific, and it increased in percent cover (>50%) up to 61 m on Bahamian reefs following the invasion of the lionfish Pterois volitans. We examined the physiological and ecological constraints contributing to the spread of Lobophora on Bahamian reefs across a mesophotic depth gradient from 30 to 61 m, pre‐ and post‐lionfish invasion. Results indicate that there were no physiological limitations to the depth distribution of Lobophora within this range prior to the lionfish invasion. Herbivory by acanthurids and scarids in algal recruitment plots at mesophotic depths was higher prior to the lionfish invasion, and Lobophora chemical defenses were ineffective against an omnivorous fish species. In contrast, Lobophora exhibited significant allelopathic activity against the coral Montastraea cavernosa and the sponge Agelas clathrodes in laboratory assays. These data indicate that when lionfish predation on herbivorous fish released Lobophora from grazing pressure at depth, Lobophora expanded its benthic cover to a depth of 61 m, where it replaced the dominant coral and sponge species. Our results suggest that this chemically defended alga may out‐compete these species in situ, and that mesophotic reefs may be further impacted in the near future as Lobophora continues to expand to its compensation point.     

The effect of the quality of food patches on larval vertical distribution of the sea urchins Lytechinus variegatus (Lamarck) and Strongylocentrotus droebachiensis (Mueller)

Although most invertebrate larvae are weak swimmers and act as passive particles on horizontal scales, they may be able to regulate their vertical position in response to different factors, including increased food concentration. We examined the effect of the quality of food patches on larval vertical distribution for the sea urchins Lytechinus variegatus and Strongylocentrotus droebachiensis, and determined the effect of dietary conditioning on that response in the laboratory. We reared larvae on a mixed algal diet of Dunaliella tertiolecta and Isochrysis galbana under low (500 cells ml⁻¹) and high (5000 cells ml⁻¹) rations. Food patches were maintained in Plexiglas rectangular columns (30×10×10 cm) using a density gradient, where practical salinity in the bottom layer was 33, in the middle layer 30, and in the top layer 27. We examined the magnitude and mechanism of a behavioural response of larvae of L. variegatus in the four-arm stage, and on two developmental stages of S. droebachiensis (four- and six-arm), by manipulating patch quality. In the absence of a patch, larvae of both species and developmental stages swam through to the surface of the experimental columns. In the presence of algae, fewer larvae were present above the patch and more were at the patch than in control columns. More larvae swam through patches of “unflavoured” algal mimics than algal patches, and aggregated at the surface. Larval distribution relative to patches of algal filtrate without algal cells or of “flavoured” algal mimics in algal filtrate was not consistently different from that in either control or algal patches; thus, the magnitude of larval response to filtrate (with or without particles) was intermediate between that to control and algal patches. For L. variegatus, more larvae crossed the patches when reared on low than high rations, indicating that poorly conditioned larvae may be less responsive to environmental cues. Our results suggest that larvae can actively aggregate and maintain a vertical position in response to a food patch that depends on the quality and quantity of food. The response appears to be based mostly on a chemosensory rather than a mechanosensory mechanism.     

Species responses of colonising beetles to variation in patch quality,number, and context in experimental aquatic landscapes

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Biotic and abiotic changes along a cyclic succession driven by shrubs in semiarid steppes from Patagonia

Aim

We studied the legacy effects of shrubs during the downgrade phase of high–cover patches. Specifically, are woody species able to modify environmental attributes at patch level to such an extent as to alter the colonization once they have vacated their original position?

Methods

We monitored five environmental variables along an experimental four-stage downgrading gradient of high–cover patches during two years in cold- and warm-seasons, individual plant growth during three years, as well as the floristic composition of patches along the same gradient after 13 years.

Results

The downgrade of high–cover patches reduces the aboveground protection due to the increase in wind speed (400–500%) and evaporation rate (43–160%) associated with shrub death and senescence. In addition, high–cover patches increase the total soil nitrogen (400–600%) and reduce the infiltration rate (44–73%) on the top layer. Leaf length and flower culms of grass tussocks were lower in bare soil patches (7.5 cm and 3) compared to whatever degradation stage of high–cover patches (9–10 cm and 18–32). Floristic composition after 13 years reveals that grass species occupied the patch stages differentially, with a disjunctive pattern among species within the Poa and Pappostipa genus.

Conclusions

Legacy effects prompted by shrubs through changes in soil properties at the horizontal plane can conditioned the patch dynamics. The ability of different plant species to cope with the spatial heterogeneity at the horizontal plane should be included as a new criterion to define plant strategies from arid ecosystems according to the gap–phase dynamics and mosaic maintenance.

     

Matrix Matters:?Differences of Grand Skink Metapopulation Parameters in Native Tussock Grasslands and Exotic Pasture Grasslands

Modelling metapopulation dynamics is a potentially very powerful tool for conservation biologists. In recent years, scientists have broadened the range of variables incorporated into metapopulation modelling from using almost exclusively habitat patch size and isolation, to the inclusion of attributes of the matrix and habitat patch quality. We investigated the influence of habitat patch and matrix characteristics on the metapopulation parameters of a highly endangered lizard species, the New Zealand endemic grand skink (Oligosoma grande) taking into account incomplete detectability. The predictive ability of the developed zxmetapopulation model was assessed through cross-validation of the data and with an independent data-set. Grand skinks occur on scattered rock-outcrops surrounded by indigenous tussock (bunch) and pasture grasslands therefore implying a metapopulation structure. We found that the type of matrix surrounding the habitat patch was equally as important as the size of habitat patch for estimating occupancy, colonisation and extinction probabilities. Additionally, the type of matrix was more important than the physical distance between habitat patches for colonisation probabilities. Detection probability differed between habitat patches in the two matrix types and between habitat patches with different attributes such as habitat patch composition and abundance of vegetation on the outcrop. The developed metapopulation models can now be used for management decisions on area protection, monitoring, and the selection of translocation sites for the grand skink. Our study showed that it is important to incorporate not only habitat patch size and distance between habitat patches, but also those matrix type and habitat patch attributes which are vital in the ecology of the target species.     

Quercus and Pinus cover are determined by landscape structure and dynamics in peri-urban Mediterranean forest patches

Successional dynamics in Mediterranean forests have been modulated by anthropogenic disturbances during thousands of years, especially in areas densely populated since ancient times. Our objective is to determine whether pine tree cover (early-successional species) and oak tree cover (late-successional species), used as a surrogate of successional stage of peri-urban fragmented forests in the Vallès lowlands (Catalonia, NE, Spain), are primarily determined by (1) climate and topography; (2) anthropogenic disturbances; (3) patch structure; or (4) patch dynamics from 1956 to 1993. Quercus spp. and Pinus spp. tree cover were separately recorded on 252 randomly selected plots of 100 m², within forest patches ranging in size from 0.25 to 218 ha. Multiple linear regressions indicated that forest patch history is the most important variable determining oak and pine tree cover: new forest patches showed higher pine and lower oak tree cover than recently split patches (i.e. those that became fragmented from large forest areas after 1956). Patches already existing as such in 1956 (pre-existent patches) showed higher pine cover than recently split patches. Oak cover increased and pine cover decreased with increasing forest connectivity of the patch. Finally, highly frequented forests were related to high cover of pines. Climatic and topographic variables were not significant. We conclude that pine and oak cover in these peri-urban forests are mainly determined by recent patch dynamics, but also by the spatial pattern of patches. However, human-induced disturbance can modulate this as there is some evidence for pine being associated with a high human frequentation.     

Spatial associations among palatable and unpalatable macroalgae: A test of associational resistance with a herbivorous amphipod

Associational resistance is the process by which plants may gain protection from spatial associations with neighbouring plants. We tested whether association with an unpalatable alga, Dictyopteris acrostichoides, affects the abundance and colonisation behaviour of the herbivorous amphipod Peramphithoe parmerong on its preferred host alga Sargassum linearifolium. Despite predictions, natural densities on S. linearifolium when surrounded by D. acrostichoides were higher than on isolated individuals of S. linearifolium. Colonisation experiments in the laboratory and the field tested the hypotheses that the observed variation in field abundance with algal neighbourhood was due to variation in the size of habitat patches, physical obstruction of host finding by D. acrostichoides and variation in the relative abundance of S. linearifolium and D. acrostichoides. None of these possible mechanisms was found to significantly alter rates of amphipod colonisation on the scales of individuals selecting among algal pieces in the laboratory or among habitat patches in the field. The failure of colonisation processes to explain observed variation in natural amphipod densities suggests that post-colonisation processes such as survival or emigration may vary with the spatial associations among algae.     

Patch colonisation in a savanna grassland

Patches (1 m diameter) were cleared in a heavily grazed and a lightly grazed savanna in South Africa; half of them were covered once with grass litter. The greatest colonisation over three years occurred on heavily grazed patches with litter, the least on open, lightly grazed patches. Annual rainfall affected colonisation rate. Basal cover was lower on patches than in the surrounding vegetation. Patch composition was weakly affected by treatment, and was partly related to the composition of adjacent vegetation, but the proportional representation of species on patches differed from the surrounding vegetation. The stoloniferous Digitaria eriantha and the obligate seed reproducers Aristida bipartita and Heteropogon contortus were major components of patch communities but were uncommon in the surrounding vegetation and in the seed bank. Setaria incrassata and Themeda triandra were the predominant components of the surrounding vegetation and of the seed bank, but Setaria established very poorly and Themeda established well only on heavily grazed patches with litter. The pattern of seedling establishment was the same as that of colonisation. Most seedlings emerged atthe beginning of the wet season, with ca. 50% mortality soon thereafter. The colonising species can be partly predicted from the availability of propagules (vegetative or seed), emergence and establishment success, and subsequent growth.     

Host–parasitoid dynamics in a fragmented landscape: Holly trees,holly leaf miners and their parasitoids

Many species inhabit fragmented landscapes, where units of resource have a patchy spatial distribution. While numerous studies have investigated how the incidence and dynamics of individual species are affected by the spatial configuration and landscape context of habitat patches, fewer studies have investigated the dynamics of multiple interacting resource and consumer species in patchy landscapes. We describe a model system for investigating host–parasitoid dynamics in a patchy landscape: a network of 166 holly trees, a specialised herbivore of holly (the leaf miner, Phytomyza ilicis (Curtis, 1948)), and its suite of parasitoids. We documented patch occupancy by P. ilicis, its density within patches, and levels of parasitism over a 6-year period, and manipulated patch occupancy by creating artificially vacant habitat patches. Essentially all patches were occupied by the herbivore in each year, suggesting that metapopulation dynamics are unlikely to occur in this system. The main determinants of densities for P. ilicis and its parasitoids were resource availability (patch size and host density, respectively). While P. ilicis is apparently not restricted by the spatial distribution of resources, densities of its parasitoids showed a weaker positive relationship with host density in more isolated patches. In patches where local extinctions were generated experimentally, P. ilicis densities and levels of parasitism recovered to pre-manipulation levels within a single generation. Furthermore, patch isolation did not significantly affect re-colonisation by hosts or parasitoids. Analysing the data at a variety of spatial scales indicates that the balance between local demography and dispersal may vary depending on the scale at which patches are defined. Taken together, our results suggest that the host and its parasitoids have dispersal abilities that exceed typical inter-patch distances. Patch dynamics are thus largely governed by dispersal rather than within-patch demography, although the role of demography is higher in larger patches.     

Dense aggregations of tube-building polychaetes: response to small-scale disturbances

Many species of tube-building polychaetes form dense aggregations or ‘patches’ within marine soft-bottom habitats. Although the ecological importance of such patches has been well studied, the effect of small-scale disturbances within patches has not been investigated. On the intertidal sandflat Drum Sands, Firth of Forth, Scotland, the tube-building spionid Pygospio elegans Claparède forms patches approximately 1-1.5 m², wherein its density is significantly greater than in surrounding sediments. This study explicitly investigates whether the initial communities colonising small-scale disturbances within P. elegans patches differ in terms of community structure and colonisation mode from those of surrounding sediments.Defaunated sediments (1/125 m²) were implanted into the sediments within and outside patches during April, August and December 1997, and sampled after 3 weeks. The results indicated that some species colonised patches and non-patches in significantly different numbers, at least at certain times of the year. Following these controlled disturbances, P. elegans larvae colonised patches in significantly higher numbers compared to non-patch areas during April 1997, but showed the opposite trend in August 1997 when larval recruitment was relatively low. Adult P. elegans colonised patch sediments in higher numbers than non-patch ones during August and December. Capitella capitata (Fabricius) consistently colonised all defaunated areas in relatively high numbers while Cerastoderma edule (L.) and Macoma balthica (L.) colonised non-patch azoic sediments in significantly higher numbers than patch azoics during April 1997. Multivariate analyses, together with nonparametric analysis of similarities (ANOSIM) tests, revealed that the community composition of the disturbed sediments was significantly different between patches and non-patches during April 1997 only. These analyses also indicated that there was a strong temporal effect on community composition since the April community was markedly different from those of August and December 1997. Sediment measurements taken during December suggested that the observed faunal differences were unlikely to have been a response to differences in bulk sediment characteristics or redox potential.It is proposed that the initial recolonisation differences were likely to have been due to the near-bed hydrodynamic effects of P. elegans tubes creating local sediment stabilisation. The ecological implications of differential recruitment following small-scale disturbances within patches compared to outside patches are discussed. The ability of P. elegans patches to enhance successful conspecific colonisation, as demonstrated here, may explain the longevity of such patches. Their persistence promotes alpha diversity and small-scale patchiness, together with providing a local source of colonists for the initial recovery of large-scale disturbances.     

Long-term dynamics of the brown macroalga <Emphasis Type="Italic">Lobophora variegata</Emphasis> on deep reefs in Curaçao

Lobophora variegata occurs in the eulittoral zone and in deep water on coral reefs in Curaçao. An analysis of the long-term (1979–2006) changes in the vertical distribution of the macroalga in permanent quadrats indicated a significant increase in cover of the deepwater community. In 1998, Lobophora covered 1 and 5% of the quadrats at 20 and 30 m, respectively. By 2006, these values had risen to 25 and 18%, precipitating a shift in abundance of corals and macroalgae at both depths. This increase coincided with losses in coral cover, possibly linked to bleaching, disease and storm-related mortality in deep water plating Agaricia corals. In contrast, macroalgae remained relatively rare (<6% cover) on shallower (10 m) and deeper (40 m) reefs despite declines in coral cover also occurring at these depths, illustrating the depth-dependent dynamics of coral reefs. Several hypotheses are suggested to explain these changes.     

The effect of territorial adult pomacentrid fishes on the recruitment and survival of juveniles on patches of coral rubble

Pomacentrus wardi Whitley is one species of a guild of territorial herbivorous pomacentrid fishes which compete for space within patches of rubble substrata on reefs of the Great Barrier. An experiment has been made on a series of adjacent patches of rubble to assess the importance of the presence of adult residents on the recruitment and survival of juvenile P. wardi, and on the use of the rubble patch by non-territorial benthic-feeding fishes. Recruitment of juvenile P. wardi, and survival of experimentally added juvenile P. wardi were both greater in patches from which all adult pomacentrids had been removed than in patches with undisturbed adult populations. Patches in which the removal of adults was only partially successful did not show these effects. Experimentally added juveniles survived only poorly, whether in undisturbed patches or in patches first denuded of adults. Denuded patches were visited by greater numbers of roving benthic-feeding fishes than were undisturbed patches. Numbers of adult pomacentrids varied little over a year in undisturbed patches, and recovered within six months in denuded patches. The results are discussed with reference to the coexistence of P. wardi and other members of its guild.     

The effect of accidental burning on habitat conditions and species composition of Molinion caeruleae meadows

This study aimed to assess the effects of accidental burning on plant cover and habitat conditions in abandoned Molinion caeruleae meadows. Field studies were conducted in Kraków (Southern Poland) in the years 2017–2019 in two line transects, each with 10 permanent plots established in permanent study plots placed in accidentally burned and unburned patches. In the burned patch, the number of species, as well as Shannon-Wiener, Pielou and Simpson indicators achieved lower values. Moreover, the cover of numerous taxa affiliated with Molinion caeruleae changed remarkably, while taxa representing the alliances Calthion and Filipendulion disappeared. The considerable dominance of hemicryptophytes was observed in burned site, while in unburned place hemicryptophytes and geophytes were represented most abundantly. In the burned patch, anemochorous and autochorous species as well as small-seeded taxa dominated, while in the unburned patch autochorous species and medium-seeded taxa prevailed. The values of Ellenberg indicators in the burned patch achieved greater (light, soil reaction), lower (nitrogen, humidity) or similar (temperature, continentality) values to the unburned patch. The majority of soil properties in the burned and unburned patches were similar with the exception of the significantly greater content of potassium and lower content of magnesium in exchangeable form noticed in the burned patch. The performed investigations showed that, although the accidental fire activity inhibited the secondary succession in Molinion caeruleae meadows, burning cannot replace traditional mowing due to the reduction of species richness and disappearance of characteristic taxa.     

Life on the edge: reproductive mode and rate of invasive <Emphasis Type="Italic">Phragmites australis</Emphasis> patch expansion

The dynamics of plant invasions from initial colonization through patch expansion are driven in part by mode of reproduction, i.e., sexual (seed) and asexual (clonal fragments and expansion) means. Expansion of existing patches—both rate and mode of spread into a matrix of varying conditions—is important for predicting potential invader impacts. In this study, we used fine-scale genetic assessments and remote sensing to describe both the rate and mode of expansion for 20 Phragmites australis patches in flooded and unflooded wetland units on the Great Salt Lake, UT. We found that the majority of Phragmites patch expansion occurred via clonal spread but we also documented instances of (potentially episodic) seedling recruitment. The mode of patch expansion, inferred from patch edge genet richness, was unrelated to flooding in the wetland unit in the preceding growing season. The rate of Phragmites patch expansion varied from 0.09 to 0.35 year^?1 and was unrelated to the mode of spread. In six patches monitored across two years, monoclonal patches stayed monoclonal, whereas patches with higher genet richness had a marked increase in diversity in the second year. The findings of the present study suggest how this partially clonal species can exploit the benefits of both sexual (i.e., genetic recombination, widespread dispersal, colonization of new areas) and asexual reproduction (i.e., stability of established clones suited to local environmental conditions) to become one of the most successful wetland plant invaders. To control this species, both forms of reproduction need to be fully addressed through targeted management actions.     

Biological activities associated to the chemodiversity of the brown algae belonging to genus <Emphasis Type="Italic">Lobophora</Emphasis> (Dictyotales,Phaeophyceae)

Although Lobophora belongs to a marine algal family (Dictyotaceae) that produces a large array of secondary metabolites, it has received little attention compared to other genera, such as Dictyota, in terms of natural compounds isolation and characterization. However, metabolites produced by Lobophora species have been found to exhibit a wide array of bioactivities including pharmacological (e.g. antibacterial, antiviral, antioxidant, antitumoral), pesticidal, and ecological. This review aims to report the state-of-the-art of the natural products isolated from Lobophora species (Dictyotales, Phaeophyceae) and their associated bioactivities. All bioactivities documented in the literature are reported, therefore including studies for which pure active substances were described, as well as studies limited to extracts or fractions. From the early 1980s until today, 49 scientific works have been published on Lobophora chemistry and bioactivity, among which 40 have reported bioactivities. Only six studies, however, have identified, characterized and tested no less than 23 bioactive pure compounds (three C₂₁ polyunsaturated alcohols, three fatty-acids, a macrolactone, 11 polyketides, a few sulfated polysaccharides, three sulfolipids, a tocopherol derivative). The present review intends to raise awareness of chemists and biologists given the recent significant taxonomic progress of this brown algal genus, which holds a promising plethora of natural products yet to be discovered with ecological and pharmacological properties.     

Spatial and temporal variation in patch occupancy and population density in a model system of an arctic Collembola species assemblage

We employed an experimental model system to investigate the mechanisms underlying patterns of patch occupancy and population density in a high arctic assemblage of Collembola species inhabiting a sedge tussock landscape on Svalbard. The replicate model systems consisted of 5 cores of the tussocks (habitat patches) imbedded in a barren matrix. Four of the patches were open so that animals could migrate between them, while there was one closed patch per system to test the effect of migration on extinction rate. There were model systems of two types: one with long and one with short inter‐patch distances to test the effect of patch isolation on colonisation and extinction rates. Each of the four most common collembolan species at the field site were introduced to two open patches per system (source patches), with the other two functioning as colonisation patches for the species. The experiment was run in an ecotrone over three identical, simulated arctic summers separated by winters of 3 weeks. Six replicates of systems with short and long inter‐patch distances were sampled at the end of each summer. The species varied markedly in their performance in both open arenas and closed patches, indicating differential responses to patch humidity, consistent with their differential distribution along the moisture gradient in the field site. The extinction – colonisation dynamics differed markedly between species as predicted from our field studies. This could partly be ascribed to differential dispersal and colonisation ability, but also to different tolerance to spatially variable patch quality and/or tendency for aggregative behaviour. Three of the species exhibited dynamics that superficially resemble what could be expected from classical metapopulation dynamics. However, there was a striking discrepancy between what would be expected from the effect of migration on the extinction rate of isolated patches (in particular closed patches) and the observed rates. Thus, metapopulation processes, such as stochastic colonisation and extinction events due to demographic stochasticity, were relatively unimportant compared to other sources of spatial variability among which subtle differences in patch quality are probably most important. We discuss the value of combining field studies with model system experiments, in particular when habitat quality cannot easily be measured in the field. However, our field and laboratory studies also emphasise the need for a thorough knowledge of species‐specific life history traits for making biologically sound interpretations based on both observational and experimental data.     

Vegetation patch dynamics in rangelands: How feedbacks between large herbivores,vegetation and soil fauna alter patches over space and through time

Aim

Large herbivore grazing is a popular conservation management tool to promote vegetation structural diversity of rangelands. However, vegetation patch dynamics, that is, how patches of grazing-defended tall vegetation and grazer-preferred short lawns shift over space and time, is poorly understood. Here, we describe a new conceptual framework for patch dynamics within rangelands, combining theories of classical cyclical succession, self-organization and multitrophic feedbacks between grazers, vegetation and bioturbating soil fauna.

Location

We use the cattle-grazed salt marsh of the island Schiermonnikoog, The Netherlands, as a model system. The grazed salt marsh is characterized by distinct tall vegetation patches dominated by the grazing-defended rush Juncus maritimus and grazing-intolerant grass Elytrigia atherica, surrounded by a matrix of grazing lawn (dominated by Festuca rubra).

The Framework

Based on previous observational and experimental studies, we propose a cyclical patch dynamic where plant species composition and structure transitions through four phases: patch initiation (a) occurs when the grazing-defended rush J. maritimus establishes in the grazed lawn. Patch establishment (b) follows when the grazing-intolerant grass E. atherica establishes in the patch due to associational defence by J. maritimus and produces a large amount of litter that attracts the key bioturbating amphipod Orchestia gammarellus. Patch expansion (c) occurs when O. gammarellus activities improve soil properties of the patch, which favours E. atherica growth, leading to E. atherica competitively displacing J. maritimus in the centre of the patch. Patch degeneration (d) follows when cattle enter the enlarged patch to consume E. atherica in the centre, trample the soil, displace O. gammarellus and decrease vegetation cover, opening space for grazing-lawn species to invade. The cycle restarts when remnants of the rush J. maritimus in the degenerated patches (or individuals recently established from seed dispersal) initiate new patches in the grazing lawn.

Synthesis

Our proposed patch-dynamic model provides a means to describe the mechanisms driving vegetation patch dynamics and serves as a foundation for further experimental and observational exploration, not only for this specific system, but more generally for grazed systems worldwide that show patches of typical grazing-defended and grazer-preferred vegetation.