Fungal endophyte‐infected leaf litter alters in‐stream microbial communities and negatively influences aquatic fungal sporulation

Endophytes are ubiquitous plant‐associated microbes and although they have the potential to alter the decomposition of infected leaf litter, this has not been well‐studied. The endophyte Rhytisma punctatum infects the leaves of Acer macrophyllum (bigleaf maple), causing the appearance of black ‘tar spots’ that persist in senesced leaves. Other foliar fungi also cause visible damage in healthy tissues of this host plant system including an unidentified bullseye‐shaped lesion, common in western Washington. Using three treatments of endophyte infection status in leaf tissue (R. punctatum‐infected, bullseye‐infected, lesion‐free), leaf litter discs were submerged in a third‐order temperate stream using mesh litter bags and harvested periodically over two months to determine the effects of litter treatment and incubation time on litter mass loss, fungal sporulation, and microbial community colonization. Litter containing symptomatic endophyte infections (Rhytisma or bullseye) had reduced sporulation of aquatic hyphomycetes, but decomposed significantly faster than lesion‐free or bullseye‐infected litter. Using amplicon‐based sequencing, we found a significant difference in bacterial communities colonizing Rhytisma‐infected and bullseye‐infected leaf litter, a significant difference in fungal communities colonizing Rhytisma‐infected leaf litter compared to the two other treatments, and a change in both community structure and relative abundances of bacterial and fungal taxa throughout the study period. Indicator Species Analysis clarified the drivers of these community shifts at the genus level. Our results show that endophyte‐associated, in‐stream sporulation and microbial community effects are observable within one species of leaf litter.     

Trubs,but no trianas: filled and empty regions of angiosperm stem length‐diameter‐mechanics space

Discrete plant habit categories such as ‘tree’, ‘shrub’, and ‘liana’ belie continuous variation in nature. To study the evolution of this continuous variation, we gathered data on stem length, diameter and tissue mechanical stiffness across a highly morphologically diverse highland xerophytic scrub on a lava flow in central Mexico. With stem allometric and mechanical data from 1216 segments from 50 species, we examined relationships between stem length–diameter proportions and tissue mechanical stiffness using linear mixed‐effects models. Rather than a series of discrete clouds in stem length–diameter–tissue stiffness space, corresponding to traditional habit categories, the plants of this xerophytic scrub formed a single continuous one. Within this cloud, self‐supporting plants had stems that became predictably longer and tissues that became stiffer for a given diameter increase, and there was no paucity of intermediates between trees and shrubs (‘trubs’). Non self‐supporting plants had a steeper stem length–diameter slope and their tissues did not increase in stiffness with stem size. The area between self‐ and non self‐supporting plants was sparsely occupied as stem size increased. We predict that this ‘empty’ space between lianas and trees is developmentally accessible but of low fitness, meaning that there should be few ‘trianas’ in nature. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 361–373.     

Non‐trophic effects of litter reduce ant predation and determine caterpillar survival and distribution

The tritrophic model featuring plants consumed by herbivores consumed by parasitoids or predators has become the primary paradigm used to describe herbivore dynamics. However, interactions involving herbivores can be habitat‐ specific and plants often provide habitat, as well as food. Structural complexity of the habitat may favor predators or may allow herbivore prey to escape detection and capture. This study considered the spatial and temporal dynamics of an arctiid caterpillar, Platyprepia virginalis. The tritrophic model that includes only a tachinid parasitoid that attacks P. virginalis and the caterpillars’ primary host‐plant, Lupinus arboreus, has failed to provide much insight into this system. Instead, we found that ants killed and removed many small caterpillars. Protecting caterpillars from ants increased their survival three‐fold and five‐fold in assays conducted during two years. Caterpillars were more likely to survive in short‐term assays at sites that naturally had a deeper cover of dead and living plant material. Experiments with baits showed that ant recruitment declined as litter depth increased on average. These survey results indicated that ant predation was an important source of mortality for young caterpillars and that the presence of thick litter reduced this mortality. These results were corroborated in an experiment that manipulated litter depth and ant access to caterpillars. Previous findings that other defoliating caterpillars increased litter depth and benefitted P. virginalis are also consistent with this hypothesis. Litter acts as an important non‐trophic resource, allowing caterpillars to avoid predation by ants such that wet sites with deep litter act as source populations for caterpillars. Our results show strong effects of both trophic and non‐trophic interactions since plants indirectly provided limiting habitat and this heterogeneous habitat strongly affected risk of predation and ultimately caterpillar abundance and distribution.     

Fungal colonization of the rooting system of the early land plant Asteroxylon mackiei from the 407‐Myr‐old Rhynie Chert (Scotland,UK)

Associations between plants and fungi were an important and varied feature of early terrestrial ecosystems, but in most instances their biological functions remain poorly understood. We document a new species of fungus colonizing the rooting system of the early lycopod Asteroxylon mackiei, based on exceptionally well‐preserved fossils from the Rhynie Chert. We investigated historical petrographic thin sections using standard optical microscopy and confocal laser scanning microscopy. P alaeozoosporites renaultii gen. nov., sp. nov. colonized the inner cortex of the plant rooting system. The fungus had an aseptate thallus with isotomous or sympodial branching. The mycelium bore distinctive porate, globose to elongated structures that we interpret as zoosporangia and resting sporangia. Doubts remain over the precise systematic affinity of P. renaultii, which closely resembles chytrids. Whereas most of the Rhynie Chert plants developed symbiotic associations of the mycorrhizal type, it seems that this was not the case for Asteroxylon mackiei, which possessed the most evolved rooting system among the Rhynie Chert plants. We argue that the new root‐borne fungus was probably parasitic. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 201–213.     

Murderous plants: Victorian Gothic,Darwin and modern insights into vegetable carnivory

Darwin's interest in carnivorous plants was in keeping with the Victorian fascination with Gothic horrors, and his experiments on them were many and varied, ranging from what appears to be idle curiosity (e.g. what will happen if I place a human hair on a Drosera leaf?) to detailed investigations of mechanisms. Mechanisms for capture and digestion of prey vary greatly among the six (or more) lineages of flowering plants that have well‐developed carnivory, and some are much more active than others. Passive carnivory is common in some groups, and one, Roridula (Roridulaceae) from southern Africa, is so passively carnivorous that it requires the presence of an insect intermediate to derive any benefit from prey trapped on its leaves. Other groups not generally considered to be carnivores, such as Stylidium (Stylidiaceae), some species of Potentilla (Rosaceae), Proboscidea (Martyniaceae) and Geranium (Geraniaceae), that have been demonstrated to both produce digestive enzymes on their epidermal surfaces and be capable of absorbing the products, are putatively just as ‘carnivorous’ as Roridula. There is no clear way to discriminate between cases of passive and active carnivory and between non‐carnivorous and carnivorous plants – all intermediates exist. Here, we document the various angiosperm clades in which carnivory has evolved and the degree to which these plants have become ‘complete carnivores’. We also discuss the problems with definition of the terms used to describe carnivorous plants. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 161 , 329–356.     

Away‐field advantage: mangrove seedlings grow best in litter from other mangrove species

Plant community composition can impact ecosystem processes via litter feedbacks. Species variation in litter quality may generate different patterns of nutrient supply for plants that are dependent on litter inputs. However, it is not known whether plants grow faster in their own litter, litter from other species, or in litter mixtures from multiple species. To test whether litter identity and mixture status influenced mangrove seedling growth, biomass allocation, and stoichiometry, we performed mesocosm experiments. Two species of mangrove seedlings, Avicennia germinans, black mangrove and Rhizophora mangle, red mangrove, were exposed to all possible combinations of three mangrove litter types and were isolated from all other nutrient inputs. Litter treatments significantly altered seedling growth. Seedlings from both mangrove species grew most rapidly in litter from a different species rather than their own, irrespective of litter chemical quality, decomposition rate, and nitrogen release. Litter mixtures from white and black mangroves caused black mangroves to grow 65% more than expected. Litter treatments did not impact seedling root:shoot ratios or tissue C:N. Our finding that seedlings grow best in litter from other species may indicate a mechanism that helps sustain the coexistence of dominant species.     

Staminal hairs enhance fecundity in the pollen‐rewarding self‐incompatible lily Bulbine abyssinica

Flowering plants typically use floral rewards to attract animal pollinators. Unlike nectar, pollen rewards are usually visible and may thus function as a signal that influences landing decisions by pollen‐seeking insects. Here we artificially manipulate the presence of both pollen and staminal hairs (a putative false signal of pollen reward availability) in the hermaphroditic lily Bulbine abyssinica (Xanthorrhoeaceae) to investigate their effects on bee visitation and fecundity, and also test for trade‐offs between pollen production and seed production. Honeybees, the primary floral visitors, are probably not able to distinguish between colours of petals, staminal hairs and pollen of B. abyssinica, according to analysis of reflectance spectra in a bee vision model. Flowers with both pollen and hairs removed had the lowest levels of bee visitation, seed set and seed abortions. Flowers containing hairs had an ～50% increase in visitation rate and seed set compared with emasculated flowers, while intact controls had the highest seed abortion rate. Ovule discounting in intact flowers is probably due to ovarian self‐incompatibility (or strong early inbreeding depression) as ovules penetrated by tubes from self‐pollen uniformly failed to develop into seeds. These results show that staminal hairs can enhance plant fecundity by increasing attraction of pollen‐seeking insects to flowers without increasing the risk of ovule discounting through pollinator‐mediated self‐pollination. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 481–490.     

Once a land of big wild rivers: specialism is context‐dependent for riparian snails (Pulmonata: Valloniidae) in central Europe

Specialist species may be perceived as such because of their narrow ecological requirements, but this may be context‐dependent. The genus Vallonia (Gastropoda Pulmonata: Valloniidae) includes widespread generalist species and also two specialists endemic to Central Europe: Vallonia suevica, restricted to warm, wet meadows and riverbanks subject to seasonal flooding; and Vallonia declivis, living only in wet to humid meadows, riverbanks, and reedbeds. Both have experienced dramatic declines; as is the case with many land snails, their global conservation status has been underestimated: these species are Critically Endangered. Other congenerics are probably dispersed by birds. In contrast, the distributions of these meadow specialists appear to be the outcome of their strictly riparian habitat coupled with dispersal by fish. Thus, they have tracked drainage changes through the Pleistocene from their origin in the floodplain uplands of the Danube biodiversity hotspot in the Pliocene. Natural dispersal mechanisms have been disrupted, and riparian and river ecosystems have been destroyed throughout Europe. This has led big‐river specialist molluscs and their associated fishes to the brink of extinction. The notion of specialism thus depends on the ecological context; it is useful to stress their non‐invasive character, current restriction to scarce habitats, and evolution under quite different conditions. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●● , ●●–●●.     

Permanent spore dyads are not ‘a thing of the past’: on their occurrence in the liverwort Haplomitrium (Haplomitriopsida)

The liverwort Haplomitrium gibbsiae is shown to regularly produce spores released in the form of permanent dyad pairs. Developmental studies indicate that the dyads are produced via a unique half‐lobed configuration of the developing sporocyte. Many fossil cryptophytes of Siluro‐Devonian age, which are clearly embryophytes based on their morphology, contain permanent spore dyads in their sporangia, but this is the first demonstration of their occurrence in a living plant, a species belonging to Haplomitriopsida, which resolves in a clade that is considered to be sister to all remaining liverworts. Dispersed spore‐like dyads are found in the rock record as far back as the mid‐Cambrian, but most researchers still regard the first occurrence of isomorphic, tetrahedral tetrads in the mid‐Ordovician as the benchmark age for the origin of land plants. Regardless of the geological antiquity of the embryophytes, it appears that H. gibbsiae has retained a non‐simultaneous form of sporogenesis that may ultimately be traced to a charophytic origin. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 658–669.     

Gone with the wind: trembling leaves may deter herbivory

Plants employ various defensive tactics against herbivores but are rarely considered to use rapid movements to resist predation. However, the aboveground parts of plants are often forcefully moved by wind and rain. This passive movement has been overlooked as an anti‐herbivore trait. The leaves of many plant species, such as aspens, Indian sacred fig, bamboos, and palms, tremble even in a slight breeze. Leaves that are easily moved by gentle winds can sometimes resist strong winds and may have other benefits as well. In the present study, it is proposed that the movement of such plant leaves physically deters arthropod herbivory and pathogen infection by repelling colonization and oviposition by herbivorous insects. This leads to herbivores and pathogens being dislodged from the plants, and the ensuing death of the herbivores on the ground or at least their recolonization to other plants, as well as the interruption of feeding, intraspecific communication and the mating behaviour of herbivores, thus lowering their performance on the plant or increasing enemy attack of the herbivores. In addition, passive leaf movements may undermine herbivore camouflage and expose them to predation, and may also allow plant volatiles to diffuse efficiently to repel herbivores and attract natural enemies. Thus, the mechanistic properties of these leaves may have anti‐herbivore effects in the wind and rain. This hypothesis can also be applied to aquatic plants that tremble in gentle water currents. In addition, genetic manipulation of the tendency for leaf movement may be beneficial for the management of pest insects and pathogens with reduced pesticides in forestry and agriculture. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 738–747.     

Gene dispersal inference across forest patches in an endangered medicinal tree: comparison of model‐based approaches

Propagule dispersal in plants is a fundamental mechanism for colonizing new sites and adapting to changing climates, as well as for maintaining genetic diversity. Contrasting past and current gene dispersal can provide useful insights to gauge the extent of recent human disturbances and guide management strategies. However, research on gene dispersal of plants is not yet exhaustive because evolutionary or environmental impacts are often species‐specific and most existing studies have focused on analysis of dispersal at a single site, which may not be helpful for landscape‐level inferences and management interventions. In the present study, we assessed whether current gene or propagule dispersal would be more restricted than past gene dispersal at multiple patches of the endangered medicinal tree, Prunus africana. We employed eight highly polymorphic microsatellite markers in conjunction with isolation‐by‐distance, spatial genetic structure (SGS), and parentage assignment models to estimate gene dispersal distance in a spatial extent of approximately 400 km². There was no significant difference between gene dispersal distances across the different models (Friedman chi‐squared = 7.286, d.f. = 5, P = 0.2002). Estimates of current gene dispersal distance were comparable to dispersal in the last few generations. However, gene dispersal distance was much shorter in smaller than bigger forest patches. Further, significant (P < 0.05) SGS was detected in most forest patches, with the extent of SGS among adults being stronger in the smaller than bigger patches. These results suggest the need for practicing enrichment plantings in most forest patches, particularly in the smaller ones, to assist gene exchange among individuals and patches. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 887–904.     

Convergent evolution of sunbird pollination systems of Impatiens species in tropical Africa and hummingbird systems of the New World

The bird pollination systems of the New and Old Worlds evolved independently, and differ in many aspects. New World plants are often presented as those adapted to hovering birds while Old World plants to perching birds. Most Neotropical studies also demonstrate that in hummingbird species rich assemblages, only a small number of highly specialized birds exploits the most specialized plants with long corollas. Nevertheless, recent research on bird–plant pollination interactions suggest that sunbird pollination systems in the Old World have converged more with the highly specialized hummingbird pollination systems than previously thought. In this study we focus on the pollination systems of the bird pollination syndrome Impatiens species on Mt. Cameroon, West Africa. We show that despite the high diversity of sunbirds on Mt. Cameroon, only Cyanomitra oritis appear to be important pollinator of all Impatiens species. This asymmetry indicates the absence of pair wise co‐evolution and points to a diffuse co‐evolutionary process resulting in guilds of highly specialized plants and birds; a situation well known from hummingbirds and specialized plant communities of the New World. Additionally, the herbaceous habits of Impatiens species, the frequent adaptations to pollination by hovering birds, and the habitat preference for understory in tropical forests or epiphytic growth, resemble the highly specialized Neotropical plants. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 127–133.     

Microhabitat associations of terrestrial insectivorous birds in Amazonian rainforest and second‐growth forests

ABSTRACT Across the Neotropics, small‐bodied terrestrial insectivores are sensitive to forest fragmentation and are largely absent from second‐growth forests. Despite their sensitivity to forest structure, the microhabitat relationships of these birds have not been quantified. From July 1994 to January 1995 in central Amazonia, we characterized habitat at sites where nine species of terrestrial insectivores were observed foraging, as well as at randomly selected sites in continuous forest and two types of 10–15‐yr‐old second‐growth forest common in Amazonia (Vismia‐ and Cecropia‐dominated). We used factor analysis to find suites of correlated variables. From each factor, we selected a representative variable that was relatively easy to measure. We used Bayesian analysis to estimate means and standard deviations of these variables for each species and for each type of habitat. All nine focal species were associated with ranges of microhabitat variables, such as leaf litter depth and tree densities, often absent in second‐growth forests. At least in the early stages of regeneration, neither type of second‐growth forest provides suitable structure for the terrestrial insectivores in our study. The large leaves of Cecropia trees that make up the thick leaf litter may preclude the use of Cecropia‐dominated second growth by our focal species, many of which manipulate leaves when foraging. The leaf litter in Vismia‐dominated second growth was also thicker than sites used for foraging by our focal species. In addition, Vismia‐dominated growth had more small trees and small nonwoody vegetation, perhaps impeding movement by terrestrial birds. In continuous forest, our focal species foraged in microhabitats with characteristics that generally overlapped those of randomly selected sites. Thus, our results are consistent with the hypothesis that microhabitat differences make second‐growth forests unsuitable for our focal species.     

Genetic diversity,sexual condition,and microhabitat preference determine mating patterns in Sphagnum (Sphagnaceae) peat‐mosses

In bryophytes, the possibility of intragametophytic selfing creates complex mating patterns that are not possible in seed plants, although relatively little is known about patterns of inbreeding in natural populations. In the peat‐moss genus Sphagnum, taxa are generally bisexual (gametophytes produce both sperm and egg) or unisexual (gametes produced by separate male and female plants). We sampled populations of 14 species, aiming to assess inbreeding variation and inbreeding depression in sporophytes, and to evaluate correlations between sexual expression, mating systems, and microhabitat preferences. We sampled maternal gametophytes and their attached sporophytes at 12–19 microsatellite loci. Bisexual species exhibited higher levels of inbreeding than unisexual species but did generally engage in some outcrossing. Inbreeding depression did not appear to be common in either unisexual or bisexual species. Genetic diversity was higher in populations of unisexual species compared to populations of bisexual species. We found a significant association between species microhabitat preference and population genetic diversity: species preferring hummocks (high above water table) had populations with lower diversity than species inhabiting hollows (at the water table). We also found a significant interaction between sexual condition, microhabitat preference, and inbreeding coefficients, suggesting a vital role for species ecology in determining mating patterns in Sphagnum populations. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 96–113.     

A three‐marker DNA barcoding approach for ecological studies of xerothermic plants and herbivorous insects from central Europe

The DNA barcoding technique developed for species identification has recently been adapted for ecological studies (e.g. host plant identification). Comprehensive barcode databases, covering most species inhabiting areas, habitats or communities of interest are essential for reliable and efficient identification of plants. Here we present a three‐barcode (plastid rbcL and matK genes and the trnL intron) database for xerothermic plant species from central Europe. About 85% of the xerothermic plant species (126 out of c. 150) known to be associated with xerothermic habitats were collected and barcoded. The database contains barcodes for 117 (rbcL and trnL) and 96 (matK) species. Interspecific nucleotide distances were in the ranges 0–17.9% (0–3.2% within genera) for rbcL, 0–44.4% (0–3.1%) for trnL and 0–52.5% (0–10.9%) for matK. Blast‐searching of each sequence in the database against the entire database showed that species‐level identification is possible for 89.6% (rbcL), 98.4% (trnL) and 96.4% (matK) of examined plant species. The utility of the presented database for identification of host plants was demonstrated using two insect species associated with xerothermic habitats: the oligophagous leaf‐beetle Cheilotoma musciformis (for which two host plants in Fabaceae were identified) and the polyphagous weevil Polydrusus inustus (which was found to feed on 14 host plants, mostly Rosaceae, Asteraceae and Fabaceae). The developed database will be useful in various applications, including biodiversity, phylogeography, conservation and ecology. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 576–592.     

Aphids decelerate litter nitrogen mineralisation through changes in litter quality

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Phylogeography of the whelk genus Cominella (Gastropoda: Buccinidae) suggests long‐distance counter‐current dispersal of a direct developer

The gastropod genus Cominella Gray, 1850 consists of approximately 20 species that inhabit a wide range of marine environments in New Zealand and Australia, including its external territory, the geographically isolated Norfolk Island. This distribution is puzzling, however, with apparently closely‐related species occurring either side of the Tasman Sea, even though all species are considered to have limited dispersal abilities. To determine how Cominella attained its current distribution, we derived a dated molecular phylogeny, which revealed a clade comprising all the Australian and Norfolk Island species nested within four clades of solely New Zealand species. This Australian clade diverged well after the vicariant separation of New Zealand from Australia, and implies two long‐distance dispersal events: a counter‐current movement across the Tasman Sea from New Zealand to Australia, occurring at the origination of the clade, followed by the colonization of Norfolk Island. The biology of Cominella suggests that the most likely method of long‐distance dispersal is rafting as egg capsules. Our robust phylogeny also means that the current Cominella classification requires revision. We propose that our clades be recognized as subgenera: Cominella (s.s.), Cominista, Josepha, Cominula, and Eucominia, with each subgenus comprising only of New Zealand or Australian species. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 315–332.     

Unmasking cryptic species: morphometric and phylogenetic analyses of the Ibero‐North African Linaria incarnata complex

Linaria incarnata has been treated as a widely distributed Ibero‐North African species in the latest taxonomic reviews. Morphological and phylogenetic disparity between populations of this species has been previously reported. Here we present new morphological and phylogenetic evidence for the disintegration of L. incarnata into three distinct species: L. incarnata from the western Iberian Peninsula; L . mamorensis sp. nov. from north‐western Morocco; and L. onubensis from south‐western Spain. The relatively poor morphological differentiation between these taxa (which can be regarded as cryptic species) and their distinct phylogenetic positions indicate that characters of the L. incarnata morphotype have been acquired multiple times in the evolution of Linaria section Versicolores. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 00, 000–000.     

Evolution of life‐history traits in geographically isolated populations of Vaejovis scorpions (Scorpiones: Vaejovidae)

Geographical isolation can over time accumulate life‐history variation which can eventually lead to speciation. We used five species of Vaejovis scorpions that have been isolated from one another since the Pleistocene glaciation to identify if biogeographical patterns have allowed for the accumulation of life‐history variation among species. Gravid females were captured and brought back to the lab until giving birth. Once offspring had begun to disperse, measurements of female size, reproductive investment, offspring size, offspring number, and variation in offspring size were recorded. Differences in how each species allocated energy to these variables were analysed utilizing path analysis and structural equation modelling. Female and offspring size, litter size, and total litter mass differed among species, but relative energetic investment did not. Most significant differences among species were not present after removing the effect of female size, indicating that female size is a major source of life‐history variation. Path analyses indicated that there was no size–number trade‐off within any species and that each species allocates energy toward total litter mass differently. Additionally, as offspring size increased, the variation in offspring mass decreased. These results show that each species allocates the same relative amount of energy in different ways. The variation seen could be a response to environmental variability or uncertainty, a product of maternal effects, or caused by the sufficient accumulation of genetic differences due to geographical isolation. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 715–727.     

The effect of plant litter on ecosystem properties in a Mediterranean semi‐arid shrubland

Abstract. We studied the distribution of litter in a shrubland of the Negev with a semi‐arid Mediterranean climate of less than 200 mm of rainfall per year. Our focus was on the effects of litter on properties of landscape patches relevant to ecosystem processes (water runoff and soil erosion), annual plant community responses (seedling density, biomass production and species richness), and animal activity (soil disturbance by termites). Three 60‐m transects, extending across a pair of opposing north‐ and south‐facing slopes and their drainage channel, showed that litter accumulates not only under shrubs, but to a lesser extent also on the crusted inter‐shrub open areas. We used 35 experimental units (‘cells’, 0.5m × 1 m), each containing a crust and a shrub patch. Because runoff flows from crusted patches and is intercepted by shrub patches, the latter were in the lower third of the cells. Leaf litter was added in single and double amounts providing ca. 0.5 and 1.0 cm litter depth, to either, both, or none of the patches. Litter addition significantly decreased the amount of runoff, regardless of the location and amount of litter applied. Litter on the crust increased species number and seedling density of species with low abundance. Adding a double litter layer increased annual plant biomass production, while a single amount had no effect. Litter addition to the shrub patch affected neither biomass nor species richness. Litter addition to both patches at both quantities caused a large increase in termite activity. Termites caused disturbance by disrupting the crust, which may contribute to the reduction in runoff amounts. In the open, flat crust patches, annual plant communities are limited in their productivity and species richness, as there are few structures stopping the outflow of water, soil and seeds. Litter adds such structures, but affects the plant communities only when added to litter‐free crust. Litter accumulation and its patchy distribution have large impacts on landscape patch properties affecting resource distribution, plant productivity and diversity, and animal activity. Therefore, understanding litter distribution in relation to the patchy structure of the landscape of semi‐arid shrubland should be viewed as an important component of shrubland management.