A Pleistocene legacy structures variation in modern seagrass ecosystems.

Distribution of Earth's biomes is structured by the match between climate and plant traits, which in turn shape associated communities and ecosystem processes and services. However, that climate-trait match can be disrupted by historical events, with lasting ecosystem impacts. As Earth's environment changes faster than at any time in human history, critical questions are whether and how organismal traits and ecosystems can adjust to altered conditions. We quantified the relative importance of current environmental forcing versus evolutionary history in shaping the growth form (stature and biomass) and associated community of eelgrass (Zostera marina), a widespread foundation plant of marine ecosystems along Northern Hemisphere coastlines, which experienced major shifts in distribution and genetic composition during the Pleistocene. We found that eelgrass stature and biomass retain a legacy of the Pleistocene colonization of the Atlantic from the ancestral Pacific range and of more recent within-basin bottlenecks and genetic differentiation. This evolutionary legacy in turn influences the biomass of associated algae and invertebrates that fuel coastal food webs, with effects comparable to or stronger than effects of current environmental forcing. Such historical lags in phenotypic acclimatization may constrain ecosystem adjustments to rapid anthropogenic climate change, thus altering predictions about the future functioning of ecosystems.

Are shark teeth proxies for functional traits? A framework to infer ecology from the fossil record.

Modern sharks have an evolutionary history of at least 250 million years and are known to play key roles in marine systems, from controlling prey populations to connecting habitats across oceans. These ecological roles can be quantified based on their functional traits, which are typically morphological (e.g., body size) or behavioural (e.g., feeding and diet). Nonetheless, the understanding of such roles of extinct sharks is limited due to the inherent incompleteness of their fossil record, which consists mainly of isolated teeth. As such, establishing links between tooth morphology and ecological traits in living sharks could provide a useful framework to infer sharks' ecology from the fossil record. Here, based on extant sharks from which morphological and behavioural characteristics are known, the authors assess the extent to which isolated teeth can serve as proxies for functional traits. To do so, they first review the scientific literature on extant species to evaluate the use of shark dental characters as proxies for ecology to then perform validation analyses based on an independent data set collected from museum collections. Their results reveal that 12 dental characters have been used in shark literature as proxies for three functional traits: body size, prey preference and feeding mechanism. From all dental characters identified, tooth size and cutting edge are the most widely used. Validation analyses suggest that seven dental characters - crown height, crown width, cutting edge, lateral cusplets, curvature, longitudinal outline and cross-section outline - are the best proxies for the three functional traits. In particular, tooth size (crown height and width) was found to be a reliable proxy of all three traits; the presence of serrations on the cutting edge was one of the best proxies for prey preference; and tooth shape (longitudinal outline) and the presence of lateral cusplets were among the best indicators of feeding mechanism. Overall, the authors' results suggest that in the absence of directly measurable traits in the fossil record, these seven dental characters (and different combinations of them) can be used to quantify the ecological roles of extinct sharks. This information has the potential to provide key insights into how shark functional diversity has changed through time, including their ecological responses to extinction events.

Loss of function mutations in CCDC32 cause a congenital syndrome characterized by craniofacial, cardiac and neurodevelopmental anomalies.

Despite the wide use of genomics to investigate the molecular basis of rare congenital malformations, a significant fraction of patients remains bereft of diagnosis. As part of our continuous effort to recruit and perform genomic and functional studies on such cohorts, we investigated the genetic and mechanistic cause of disease in two independent consanguineous families affected by overlapping craniofacial, cardiac, laterality and neurodevelopmental anomalies. Using whole exome sequencing, we identified homozygous frameshift CCDC32 variants in three affected individuals. Functional analysis in a zebrafish model revealed that ccdc32 depletion recapitulates the human phenotypes. Because some of the patient phenotypes overlap defects common to ciliopathies, we asked if loss of CCDC32 might contribute to the dysfunction of this organelle. Consistent with this hypothesis, we show that ccdc32 is required for normal cilia formation in zebrafish embryos and mammalian cell culture, arguing that ciliary defects are at least partially involved in the pathomechanism of this disorder.

Novel truncating mutations in CTNND1 cause a dominant craniofacial and cardiac syndrome.

CTNND1 encodes the p120-catenin (p120) protein, which has a wide range of functions, including the maintenance of cell-cell junctions, regulation of the epithelial-mesenchymal transition and transcriptional signalling. Due to advances in next-generation sequencing, CTNND1 has been implicated in human diseases including cleft palate and blepharocheilodontic (BCD) syndrome albeit only recently. In this study, we identify eight novel protein-truncating variants, six de novo, in 13 participants from nine families presenting with craniofacial dysmorphisms including cleft palate and hypodontia, as well as congenital cardiac anomalies, limb dysmorphologies and neurodevelopmental disorders. Using conditional deletions in mice as well as CRISPR/Cas9 approaches to target CTNND1 in Xenopus, we identified a subset of phenotypes that can be linked to p120-catenin in epithelial integrity and turnover, and additional phenotypes that suggest mesenchymal roles of CTNND1. We propose that CTNND1 variants have a wider developmental role than previously described and that variations in this gene underlie not only cleft palate and BCD but may be expanded to a broader velocardiofacial-like syndrome.

The biogeography of community assembly: latitude and predation drive variation in community trait distribution in a guild of epifaunal crustaceans.

While considerable evidence exists of biogeographic patterns in the intensity of species interactions, the influence of these patterns on variation in community structure is less clear. Studying how the distributions of traits in communities vary along global gradients can inform how variation in interactions and other factors contribute to the process of community assembly. Using a model selection approach on measures of trait dispersion in crustaceans associated with eelgrass (Zostera marina) spanning 30° of latitude in two oceans, we found that dispersion strongly increased with increasing predation and decreasing latitude. Ocean and epiphyte load appeared as secondary predictors; Pacific communities were more overdispersed while Atlantic communities were more clustered, and increasing epiphytes were associated with increased clustering. By examining how species interactions and environmental filters influence community structure across biogeographic regions, we demonstrate how both latitudinal variation in species interactions and historical contingency shape these responses. Community trait distributions have implications for ecosystem stability and functioning, and integrating large-scale observations of environmental filters, species interactions and traits can help us predict how communities may respond to environmental change.

RPSA, a candidate gene for isolated congenital asplenia, is required for pre-rRNA processing and spleen formation in Xenopus.

A growing number of tissue-specific inherited disorders are associated with impaired ribosome production, despite the universal requirement for ribosome function. Recently, mutations in RPSA, a protein component of the small ribosomal subunit, were discovered to underlie approximately half of all isolated congenital asplenia cases. However, the mechanisms by which mutations in this ribosome biogenesis factor lead specifically to spleen agenesis remain unknown, in part due to the lack of a suitable animal model for study. Here we reveal that RPSA is required for normal spleen development in the frog, Xenopus tropicalis Depletion of Rpsa in early embryonic development disrupts pre-rRNA processing and ribosome biogenesis, and impairs expression of the key spleen patterning genes nkx2-5, bapx1 and pod1 in the spleen anlage. Importantly, we also show that whereas injection of human RPSA mRNA can rescue both pre-rRNA processing and spleen patterning, injection of human mRNA bearing a common disease-associated mutation cannot. Together, we present the first animal model of RPSA-mediated asplenia and reveal a crucial requirement for RPSA in pre-rRNA processing and molecular patterning during early Xenopus development.

Biodiversity increases the resistance of ecosystem productivity to climate extremes.

It remains unclear whether biodiversity buffers ecosystems against climate extremes, which are becoming increasingly frequent worldwide. Early results suggested that the ecosystem productivity of diverse grassland plant communities was more resistant, changing less during drought, and more resilient, recovering more quickly after drought, than that of depauperate communities. However, subsequent experimental tests produced mixed results. Here we use data from 46 experiments that manipulated grassland plant diversity to test whether biodiversity provides resistance during and resilience after climate events. We show that biodiversity increased ecosystem resistance for a broad range of climate events, including wet or dry, moderate or extreme, and brief or prolonged events. Across all studies and climate events, the productivity of low-diversity communities with one or two species changed by approximately 50% during climate events, whereas that of high-diversity communities with 16-32 species was more resistant, changing by only approximately 25%. By a year after each climate event, ecosystem productivity had often fully recovered, or overshot, normal levels of productivity in both high- and low-diversity communities, leading to no detectable dependence of ecosystem resilience on biodiversity. Our results suggest that biodiversity mainly stabilizes ecosystem productivity, and productivity-dependent ecosystem services, by increasing resistance to climate events. Anthropogenic environmental changes that drive biodiversity loss thus seem likely to decrease ecosystem stability, and restoration of biodiversity to increase it, mainly by changing the resistance of ecosystem productivity to climate events.

Alkylglycerol monooxygenase, a heterotaxy candidate gene, regulates left-right patterning via Wnt signaling.

Congenital heart disease (CHD) is a major cause of morbidity in the pediatric population yet its genetic and molecular causes remain poorly defined. Previously, we identified AGMO as a candidate heterotaxy disease gene, a disorder of left-right (LR) patterning that can have a profound effect on cardiac function. AGMO is the only known alkylglycerol monooxygenase, an orphan tetrahydrobiopterin dependent enzyme that cleaves the ether linkage in alkylglycerols. However, whether AGMO plays a role in LR patterning was unexplored. Here we reveal that Agmo is required for correct development of the embryonic LR axis in Xenopus embryos recapitulating the patient's heterotaxy phenotype. Mechanistically, we demonstrate that Agmo is a regulator of canonical Wnt signaling, required during gastrulation for normal formation of the left - right organizer. Mutational analysis demonstrates that this function is dependent on Agmo's alkylglycerol monooxygenase activity. Together, our findings identify Agmo as a regulator of canonical Wnt signaling, demonstrate a role for Agmo in embryonic axis formation, and provide insight into the poorly understood developmental requirements for ether lipid cleavage.

Below-ground biomass of plants, with a key contribution of buried shoots, increases foredune resistance to wave swash.

BACKGROUND AND AIMS: Sand dunes reduce the impact of storms on shorelines and human infrastructure. The ability of these ecosystems to provide sustained coastal protection under persistent wave attack depends on their resistance to erosion. Although flume experiments show that roots of perennial plants contribute to foredune stabilization, the role of other plant organs, and of annual species, remains poorly studied. Furthermore, it remains unknown if restored foredunes provide the same level of erosion resistance as natural foredunes. We investigated the capacity of three widespread pioneer foredune species (the perennial Ammophila arenaria and the annuals Cakile maritima and Salsola kali) to resist dune erosion, and compared the erosion resistance of Ammophila at natural and restored sites. METHODS: Cores collected in the field were tested in a flume that simulated a wave swash. A multi-model inference approach was used to disentangle the contributions of different below-ground compartments (i.e. roots, rhizomes, buried shoots) to erosion resistance. KEY RESULTS: All three species reduced erosion, with Ammophila having the strongest effect (36 % erosion reduction versus unvegetated cores). Total below-ground biomass (roots, rhizomes and shoots), rather than any single compartment, most parsimoniously explained erosion resistance. Further analysis revealed that buried shoots had the clearest individual contribution. Despite similar levels of total below-ground biomass, coarser sediment reduced erosion resistance of Ammophila cores from the restored site relative to the natural site. CONCLUSIONS: The total below-ground biomass of both annual and perennial plants, including roots, rhizomes and buried shoots, reduced dune erosion under a swash regime. Notably, we show that (1) annual pioneer species offer erosion protection, (2) buried shoots are an important plant component in driving sediment stabilization, and (3) management must consider both biological (plants and their traits) and physical (grain size) factors when integrating dunes into schemes for coastal protection.

Functional diversity of habitat formers declines scale-dependently across an environmental stress gradient.

Marine habitat formers such as seaweeds and corals are lynchpins of coastal ecosystems, but their functional diversity and how it varies with scale and context remains poorly studied. Here, we investigate the functional diversity of seaweed assemblages across the rocky intertidal stress gradient at large (zones) and small (quadrat) scales. We quantified complementary metrics of emergent group richness, functional richness (functional space occupied) and functional dispersion (trait complementarity of dominant species). With increasing shore height, under species loss and turnover, responses of functional diversity were scale- and metric-dependent. At the large scale, functional richness contracted while-notwithstanding a decline in redundancy-emergent group richness and functional dispersion were both invariant. At the small scale, all measures declined, with the strongest responses evident for functional and emergent group richness. Comparisons of observed versus expected values based on null models revealed that functional richness and dispersion were greater than expected in the low shore but converged with expected values higher on the shore. These results show that functional diversity of assemblages of marine habitat formers can be especially responsive to environmental stress gradients at small scales and for richness measures. Furthermore, niche-based processes at the small-neighbourhood-scale can favour co-occurrence of functionally distinctive species under low, but not high, stress, magnifying differences in functional diversity across environmental gradients. As assemblages of marine habitat formers face accelerating environmental change, further studies examining multiple aspects of functional diversity are needed to elucidate patterns, processes, and ecosystem consequences of community (dis-)assembly across diverse groups.

Latitude, temperature, and habitat complexity predict predation pressure in eelgrass beds across the Northern Hemisphere.

Latitudinal gradients in species interactions are widely cited as potential causes or consequences of global patterns of biodiversity. However, mechanistic studies documenting changes in interactions across broad geographic ranges are limited. We surveyed predation intensity on common prey (live amphipods and gastropods) in communities of eelgrass (Zostera marina) at 48 sites across its Northern Hemisphere range, encompassing over 37° of latitude and four continental coastlines. Predation on amphipods declined with latitude on all coasts but declined more strongly along western ocean margins where temperature gradients are steeper. Whereas in situ water temperature at the time of the experiments was uncorrelated with predation, mean annual temperature strongly positively predicted predation, suggesting a more complex mechanism than simply increased metabolic activity at the time of predation. This large-scale biogeographic pattern was modified by local habitat characteristics; predation declined with higher shoot density both among and within sites. Predation rates on gastropods, by contrast, were uniformly low and varied little among sites. The high replication and geographic extent of our study not only provides additional evidence to support biogeographic variation in predation intensity, but also insight into the mechanisms that relate temperature and biogeographic gradients in species interactions.

The ribosome biogenesis factor Nol11 is required for optimal rDNA transcription and craniofacial development in Xenopus.

The production of ribosomes is ubiquitous and fundamental to life. As such, it is surprising that defects in ribosome biogenesis underlie a growing number of symptomatically distinct inherited disorders, collectively called ribosomopathies. We previously determined that the nucleolar protein, NOL11, is essential for optimal pre-rRNA transcription and processing in human tissue culture cells. However, the role of NOL11 in the development of a multicellular organism remains unknown. Here, we reveal a critical function for NOL11 in vertebrate ribosome biogenesis and craniofacial development. Nol11 is strongly expressed in the developing cranial neural crest (CNC) of both amphibians and mammals, and knockdown of Xenopus nol11 results in impaired pre-rRNA transcription and processing, increased apoptosis, and abnormal development of the craniofacial cartilages. Inhibition of p53 rescues this skeletal phenotype, but not the underlying ribosome biogenesis defect, demonstrating an evolutionarily conserved control mechanism through which ribosome-impaired craniofacial cells are removed. Excessive activation of this mechanism impairs craniofacial development. Together, our findings reveal a novel requirement for Nol11 in craniofacial development, present the first frog model of a ribosomopathy, and provide further insight into the clinically important relationship between specific ribosome biogenesis proteins and craniofacial cell survival.

Warming magnifies predation and reduces prey coexistence in a model litter arthropod system.

Climate warming can destabilize interactions between competitors as smaller organisms gain advantages in warmer environments. Whether and how warming-induced effects on competitive interactions are modified by predation remains unknown. We hypothesized that predation will offset the competitive advantage of smaller prey species in warmer environments because of their greater vulnerability to predation. To test this, we assembled a litter arthropod community with two Collembola species (Folsomia candida and Proisotoma minuta) of different body sizes across a temperature gradient (three thermal environments) and in the presence and absence of predatory mites. Predatory mites reduced Collembola coexistence with increasing temperatures. Contradicting our hypothesis, the larger prey species always outperformed the smaller prey species in warmer environments with predators. Larger prey probably benefited as they expressed a greater trait (body length) plasticity to warming. Warming can thus magnify predation effects and reduce the probability of prey coexistence.

Expression of ribosomopathy genes during Xenopus tropicalis embryogenesis.

BACKGROUND: Because ribosomes are ubiquitously required for protein production, it was long assumed that any inherited defect in ribosome manufacture would be embryonically lethal. However, several human congenital diseases have been found to be associated with mutations in ribosome biogenesis factors. Surprisingly, despite the global requirement for ribosomes, these "ribosomopathies" are characterized by distinct and tissue specific phenotypes. The reasons for such tissue proclivity in ribosomopathies remain mysterious but may include differential expression of ribosome biogenesis factors in distinct tissues. METHODS: Here we use in situ hybridization of labeled antisense mRNA probes and ultra high temporal resolution RNA-Seq data to examine and compare expression of 13 disease associated ribosome biogenesis factors at six key stages in Xenopus tropicalis development. RESULTS: Rather than being ubiquitously expressed during development, mRNAs of all examined ribosome biogenesis factors were highly enriched in specific tissues, including the cranial neural crest and ventral blood islands. Interestingly, expression of ribosome biogenesis factors demonstrates clear differences in timing, transcript number and tissue localization. CONCLUSION: Ribosome biogenesis factor expression is more spatiotemporally regulated during embryonic development than previously expected and correlates closely with many of the common ribosomopathy phenotypes. Our findings provide information on the dynamic use of ribosome production machinery components during development and advance our understanding of their roles in disease.

Foundation species' overlap enhances biodiversity and multifunctionality from the patch to landscape scale in southeastern United States salt marshes.

Although there is mounting evidence that biodiversity is an important and widespread driver of ecosystem multifunctionality, much of this research has focused on small-scale biodiversity manipulations. Hence, which mechanisms maintain patches of enhanced biodiversity in natural systems and if these patches elevate ecosystem multifunctionality at both local and landscape scales remain outstanding questions. In a 17 month experiment conducted within southeastern United States salt marshes, we found that patches of enhanced biodiversity and multifunctionality arise only where habitat-forming foundation species overlap--i.e. where aggregations of ribbed mussels (Geukensia demissa) form around cordgrass (Spartina alterniflora) stems. By empirically scaling up our experimental results to the marsh platform at 12 sites, we further show that mussels--despite covering only approximately 1% of the marsh surface--strongly enhance five distinct ecosystem functions, including decomposition, primary production and water infiltration rate, at the landscape scale. Thus, mussels create conditions that support the co-occurrence of high densities of functionally distinct organisms within cordgrass and, in doing so, elevate salt marsh multifunctionality from the patch to landscape scale. Collectively, these findings suggest that patterns in foundation species' overlap drive variation in biodiversity and ecosystem functioning within and across natural ecosystems.We therefore argue that foundation species should be integrated in our conceptual understanding of forces that moderate biodiversity--ecosystem functioning relationships, approaches for conserving species diversity and strategies to improve the multifunctionality of degraded ecosystems.

The sign of cascading predator effects varies with prey traits in a detrital system.

Theory and experiments show that the nature of 'green' trophic cascades, between predators, herbivores and plants, varies with several key species traits: predator hunting mode and predator and prey habitat domains. Meanwhile, 'brown' cascades between predators, environment-modifying detritivores and plants have been largely overlooked and the roles of species traits, particularly prey traits, in determining the nature of these cascades remains unclear. We hypothesize that, in predator-detritivore-plant interaction chains, the burrowing ability of plant-facilitating detritivores determines their response to predators and thus the sign of indirect effect transmitted. In the dung-decomposer food web of an alpine meadow, we predicted that in the presence of above-ground predacious beetles: (i) non-burrowing detritivores will suffer mortality due to predation and transmit negative indirect effects to plants, whereas (ii) burrowing detritivores will escape predation by retreating deeper into the soil, transmitting positive indirect effects to plants. In support of predictions, experiments showed that a single species of predacious beetle (i) reduced the density of the non-burrowing species and indirectly reduced dung loss rate, soil nutrient concentrations and plant biomass, but (ii) drove the burrowing species deeper, indirectly improved soil conditions and increased plant biomass. These results show that the burrowing ability of a detritivore can determine whether it transmits a negative indirect effect mediated by a reduction in its density, or a positive indirect effect mediated by its behavioural response to predation risk. We call for further tests of our detritivore-trait hypothesis in different regions and ecosystems to further develop a general trait-based framework for trophic cascades in detrital food webs. We further advance the general hypothesis that the locomotion traits of prey species (e.g. burrowing/flying ability) may help explain their behavioural response to predation risk and the nature of indirect effect they transmit from predators to plants.

Fgf8 dosage determines midfacial integration and polarity within the nasal and optic capsules.

Craniofacial development requires an exquisitely timed and positioned cross-talk between the embryonic cephalic epithelia and mesenchyme. This cross-talk underlies the precise translation of patterning processes and information into distinct, appropriate skeletal morphologies. The molecular and cellular dialogue includes communication via secreted signaling molecules, including Fgf8, and effectors of their interpretation. Herein, we use genetic attenuation of Fgf8 in mice and perform gain-of-function mouse-chick chimeric experiments to demonstrate that significant character states of the frontonasal and optic skeletons are dependent on Fgf8. Notably, we show that the normal orientation and polarity of the nasal capsules and their developing primordia are dependent on Fgf8. We further demonstrate that Fgf8 is required for midfacial integration, and provide evidence for a role for Fgf8 in optic capsular development. Taken together, our data highlight Fgf8 signaling in craniofacial development as a plausible target for evolutionary selective pressures.

Pattern and polarity in the development and evolution of the gnathostome jaw: both conservation and heterotopy in the branchial arches of the shark, Scyliorhinus canicula.

The acquisition of jaws constitutes a landmark event in vertebrate evolution, one that in large part potentiated their success and diversification. Jaw development and patterning involves an intricate spatiotemporal series of reciprocal inductive and responsive interactions between the cephalic epithelia and the cranial neural crest (CNC) and cephalic mesodermal mesenchyme. The coordinated regulation of these interactions is critical for both the ontogenetic registration of the jaws and the evolutionary elaboration of variable jaw morphologies and designs. Current models of jaw development and evolution have been built on molecular and cellular evidence gathered mostly in amniotes such as mice, chicks and humans, and augmented by a much smaller body of work on the zebrafish. These have been partnered by essential work attempting to understand the origins of jaws that has focused on the jawless lamprey. Chondrichthyans (cartilaginous fish) are the most distant group to amniotes within extant gnathostomes, and comprise the crucial clade uniting amniotes and agnathans; yet despite their critical phylogenetic position, evidence of the molecular and cellular underpinnings of jaw development in chondrichthyans is still lacking. Recent advances in genome and molecular developmental biology of the lesser spotted dogfish shark, Scyliorhinus canicula, make it ideal for the molecular study of chondrichthyan jaw development. Here, following the 'Hinge and Caps' model of jaw development, we have investigated evidence of heterotopic (relative changes in position) and heterochronic (relative changes in timing) shifts in gene expression, relative to amniotes, in the jaw primordia of S. canicula embryos. We demonstrate the presence of clear proximo-distal polarity in gene expression patterns in the shark embryo, thus establishing a baseline molecular baüplan for branchial arch-derived jaw development and further validating the utility of the 'Hinge and Caps' model in comparative studies of jaw development and evolution. Moreover, we correlate gene expression patterns with the absence of a lambdoidal junction (formed where the maxillary first arch meets the frontonasal processes) in chondrichthyans, further highlighting the importance of this region for the development and evolution of jaw structure in advanced gnathostomes.

Cascading effects of predator-detritivore interactions depend on environmental context in a Tibetan alpine meadow.

Studies of grazing food webs show that species traits can interact with environmental factors to determine the strength of trophic cascades, but analogous context dependencies in detrital food webs remain poorly understood. In predator-detritivore-plant interaction chains, predators are expected to indirectly suppress plant biomass by reducing the density of plant-facilitating detritivores. However, this outcome can be reversed where above-ground predators drive burrowing detritivores to lower soil levels, strengthening their plant-facilitating effects. Here, we show that these trait-mediated indirect interactions further depend on environmental context in a Tibetan alpine meadow. In our study system, undulating topography generates higher (dry soil) patches interspersed with lower (wet soil) patches. Because the ability of detritivores to form deep burrows is likely to be limited by oxygen availability in low patches (wet soil), we hypothesized that (i) burrowing detritivores would undergo a vertical habitat shift, allowing them to more effectively avoid predation, in high - but not low - patches, and (ii) this shift would transmit positive effects of predators to plants in high patches by improving conditions in the lower soil layer. We tested these hypotheses using complementary field and glasshouse experiments examining whether the cascading effects of above-ground predatory beetles (presence/absence) on the density and behaviour of tunnel-forming detritivorous beetles, soil properties, and plant growth varied with patch type (low/high). Results revealed that predatory beetles did not reduce the density of detritivores in either patch type but had context-dependent trait-mediated effects, increasing the tunnelling depth of detritivores, improving soil conditions and ultimately increasing plant biomass in the high but not low patches. This study adds to an emerging predictive framework linking predators to plants in detritus food webs, demonstrating that these indirect interactions depend not just on the relative habitat domains of predators and prey, but also on environmental conditions that can predictably constrain the behavioural response of detritivores to predation risk.

Scale-dependent topographic complexity underpins abundance and spatial distribution of ecosystem engineers on natural and artificial structures.

In response to ongoing coastal urbanization, it is critical to develop effective methods to improve the biodiversity and ecological sustainability of artificial shorelines. Enhancing the topographic complexity of coastal infrastructure through the mimicry of natural substrata may facilitate the establishment of ecosystem engineering species and associated biogenic habitat formation. However, interactions between ecosystem engineers and their substratum are likely determined by organismal size and resource needs, thus making responses to topography highly scale-dependent. Here, we assessed the topographic properties (rugosity, surface area, micro-surface orientations) that underpin the abundance and distribution of two ecosystem engineers (fucoids, limpets) across six spatial scales (1-500 mm). Furthermore, we assessed the 'biogenic' rugosity created by barnacle matrices across fine scales (1-20 mm). Field surveys and 3D scanning, conducted across natural and artificial substrata, showed major effects of rugosity and associated topographic variables on ecosystem engineer assemblages and spatial occupancy, while additional abiotic environmental factors (compass direction, wave exposure) and biotic associations only had weak influences. Natural substrata exhibited ≤67 % higher rugosity than artificial ones. Fucoid-covered patches were predominantly associated with high-rugosity substrata and horizontal micro-surfaces, while homescars of limpets (≥15 mm shell length) predominated on smoother substratum patches. Barnacle-driven rugosity homogenized substrata at scales ≤10 mm. Our findings suggest that scale-dependent rugosity is a key driver of fucoid habitat formation and limpet habitat use, with wider eco-engineering applications for mimicking ecologically impactful topography on coastal infrastructure.