Changes in the composition of invertebrate assemblages from wave-exposed intertidal mussel stands along the Nova Scotia coast, Canada.

Rocky intertidal habitats occur worldwide and are mainly characterized by primary space holders such as seaweeds and sessile invertebrates. Some of these organisms are foundation species, as they can form structurally complex stands that host many small invertebrates. The abundance of primary space holders is known to vary along coastlines driven directly or indirectly by environmental variation. However, it is less clear if the invertebrate assemblages associated to a foundation species may remain relatively unchanged along coastlines, as similar stands of a foundation species can generate similar microclimates. We examined this question using abundance data for invertebrate species found in mussel stands of a similar structure in wave-exposed rocky habitats at mid-intertidal elevations along the Atlantic coast of Nova Scotia (Canada). While the most abundant invertebrate species were found at three locations spanning 315 km of coastline, species composition (a combined measure of species identity and their relative abundance) differed significantly among the locations. One of the species explaining the highest amount of variation among locations (a barnacle) exhibited potential signs of bottom-up regulation involving pelagic food supply, suggesting benthic-pelagic coupling. The abundance of the species that explained the highest amount of variation (an oligochaete) was positively related to the abundance of their predators (mites), further suggesting bottom-up forcing in these communities. Overall, we conclude that species assemblages associated to structurally similar stands of a foundation species can show clear changes in species composition at a regional scale.

Global taxonomic and functional patterns in invertebrate assemblages from rocky-intertidal mussel beds.

Mussels form extensive beds in rocky intertidal habitats on temperate seashores worldwide. They are foundation species because their beds host many invertebrates. Mussels and their associated species differ taxonomically among biogeographic regions, but all mussel beds exhibit similar structural and functional properties. Therefore, we investigated if rocky-intertidal mussel beds from around the globe host associated communities that are functionally similar despite their underlying taxonomic differences. We gathered datasets on the abundance of invertebrates found in rocky-intertidal mussel beds from the eastern and western boundaries of the Pacific and Atlantic Oceans from both hemispheres and, then, we compared their taxonomic and functional properties. Taxonomic composition differed markedly among coasts when analyzed at the taxonomic resolution reported by the surveys (often species). However, taxonomic groups with similar ecologies (28 groups including barnacles, decapods, gastropods, polychaetes, etc.) were more universally present in mussel beds. Concomitantly, functional categories of trophic level, body type, and mobility were almost always present on all studied coasts. These taxonomic groups and trait categories, however, showed regional patterns based on their relative abundances. Overall, the ability of mussel beds to host a core community type based on taxonomic groups and functional traits emphasizes their importance for biodiversity and community functioning, making them critical organisms to preserve.

Foundation species canopies affect understory beta diversity differently depending on species mobility.

Beta diversity measures the spatial variation in species composition. Because it influences several community attributes, studies are increasingly investigating its drivers. Spatial environmental heterogeneity is a major determinant of beta diversity, but canopy-forming foundation species can locally modify environmental properties. We used intertidal communities dominated by the canopy-forming alga Mazzaella laminarioides as a model system to examine how a foundation species affects spatial environmental heterogeneity and the resulting beta diversity. Since canopies were found to reduce the spatial variation of temperature and desiccation during low tides, we hypothesized that canopies would decrease understory beta diversity, which we tested through a field experiment that contrasted canopy removal with presence treatments over 32 months. The beta diversity of sessile species was always lower under canopies, but canopies never affected the beta diversity of mobile species. The observed responses for sessile species may result from their abundance being more dependent on spatial abiotic variation than for mobile species, which can occur in stressful areas while temporarily foraging or in transit to other areas. These responses may likely apply to other systems exhibiting canopy-forming foundation species hosting sessile and mobile species assemblages.

Non-consumptive effects of a predatory snail (Acanthina monodon) on a dominant habitat-forming mussel species (Perumytilus purpuratus).

Predators can influence prey through direct consumption as well as through non-consumptive effects (NCEs). NCEs usually occur mediated by behavioral changes in the prey upon detection of predator cues. Such changes may involve reduction of feeding with a variety of physiological consequences. We evaluated NCEs from an intertidal predatory snail (Acanthina monodon) on a dominant habitat-forming mussel species (Perumytilus purpuratus) from the southeastern Pacific coast. We tested whether A. monodon exerts negative NCEs on clearance rate, oxygen consumption rate, biodeposit production, and between-valve gap size in P. purpuratus. We found that waterborne predator cues triggered a decrease in these variables except biodeposit production. However, the organic content of the biodeposits increased in the presence of predator cues. The snail's physical contact with the mussels strengthened the negative NCEs on between-valve gap size. Since P. purpuratus is a dominant filter-feeder and foundation species in rocky intertidal habitats, predator NCEs on this species might indirectly influence ecosystem-level processes and community structure.

Community-wide consequences of nonconsumptive predator effects on a foundation species.

Predators can exert nonconsumptive effects (NCEs) on prey, which often take place through prey behavioural adjustments to minimise predation risk. As NCEs are widespread in nature, interest is growing to determine whether NCEs on a prey species can indirectly influence several other species simultaneously, thus leading to changes in community structure. In this study, we investigate whether a predator can exert NCEs on a foundation species and indirectly affect community structure. Through laboratory experiments, we first tested whether the predatory marine snail Acanthina monodon exerts negative NCEs on larviphagy (consumption of pelagic larvae) and phytoplankton filtration rates of the mussel Perumytilus purpuratus, an intertidal foundation species. These hypotheses stem from the notion that mussels may decrease feeding activities in the presence of predator cues to limit detection by predators. Afterwards, a field experiment tested whether the presence of A. monodon near mussel beds leads to higher colonisation rates of invertebrates that reproduce through pelagic larvae (expected under a lower larviphagy in P. purpuratus) and to a lower algal biomass on P. purpuratus shells (expected under a lower metabolite excretion in the mussels), thereby changing the community structure of the species typically found in P. purpuratus beds. The laboratory experiments revealed that waterborne cues from A. monodon limit the larviphagy and filtration rates of P. purpuratus. In turn, the field experiment showed that A. monodon cues led to greater abundances of barnacles and bivalves and a lower algal biomass in P. purpuratus beds, thus altering community structure. Overall, this study shows that a predator can indirectly affect community structure through NCEs on an invertebrate foundation species. As invertebrate foundation species are ubiquitous worldwide, understanding predator NCEs on these organisms could help to better understand community regulation in systems structured by such species.

The BenBioDen database, a global database for meio-, macro- and megabenthic biomass and densities.

Benthic fauna refers to all fauna that live in or on the seafloor, which researchers typically divide into size classes meiobenthos (32/64 µm-0.5/1 mm), macrobenthos (250 µm-1 cm), and megabenthos (>1 cm). Benthic fauna play important roles in bioturbation activity, mineralization of organic matter, and in marine food webs. Evaluating their role in these ecosystem functions requires knowledge of their global distribution and biomass. We therefore established the BenBioDen database, the largest open-access database for marine benthic biomass and density data compiled so far. In total, it includes 11,792 georeferenced benthic biomass and 51,559 benthic density records from 384 and 600 studies, respectively. We selected all references following the procedure for systematic reviews and meta-analyses, and report biomass records as grams of wet mass, dry mass, or ash-free dry mass, or carbon per m2 and as abundance records as individuals per m2. This database provides a point of reference for future studies on the distribution and biomass of benthic fauna.

Abundance data for invertebrate assemblages from intertidal mussel beds along the Atlantic Canadian coast.

This data set describes the abundance of 50 invertebrate taxa found in intertidal mussel beds along the Atlantic Canadian coast. This information resulted from a regional-scale study that investigated the effects of wave exposure on the richness and composition of invertebrate assemblages from intertidal mussel beds. Abundance data are provided for taxa representing the Annelida, Arthropoda, Bryozoa, Chordata, Cnidaria, Echinodermata, Mollusca, Nematoda, Nemertea, and Platyhelminthes. The data characterize mussel beds from wave-sheltered and wave-exposed locations spanning 315 km of the coast of Nova Scotia. Univariate and multivariate analyses revealed that the compositional structure of these invertebrate assemblages differed markedly depending on wave exposure. Overall, because of its taxonomic diversity, the inclusion of data for basal, intermediate, and top trophic levels, and the coverage of two extremes of environmental stress, this data set could be useful to test broader aspects of ecological theory. Areas of ecology that could advance using this data set are those concerning environmental stress models of community organization, abundance-occupancy relationships, species co-occurrence, species abundance distributions, dominance and rarity, spatial scales of population and community variation, and distribution of functional and phylogenetic diversity. Use of this data set for academic or educational purposes is allowed as long as the data source is properly cited. When used for academic or educational purposes, this data set should be cited using the title of this Data Paper, the names of the authors, the year of publication, and the corresponding volume and article numbers.

A 5-year study (2014-2018) of the relationship between coastal phytoplankton abundance and intertidal barnacle size along the Atlantic Canadian coast.

Benthic-pelagic coupling refers to the ecological relationships between benthic and pelagic environments. Studying such links is particularly useful to understand biological variation in intertidal organisms along marine coasts. Filter-feeding invertebrates are ecologically important on marine rocky shores, so they have often been used to investigate benthic-pelagic coupling. Most studies, however, have been conducted on eastern ocean boundaries. To evaluate benthic-pelagic coupling on a western ocean boundary, we conducted a 5-year study spanning 415 km of the Atlantic coast of Nova Scotia (Canada). We hypothesized that the summer size of intertidal barnacles (Semibalanus balanoides) recruited in the preceding spring would be positively related to the nearshore abundance (biomass) of phytoplankton, as phytoplankton constitutes food for the nauplius larvae and benthic stages of barnacles. Every year between 2014 and 2018, we measured summer barnacle size in clearings created before spring recruitment on the rocky substrate at eight wave-exposed locations along this coast. We then examined the annual relationships between barnacle size and chlorophyll-a concentration (Chl-a), a proxy for phytoplankton biomass. For every year and location, we used satellite data to calculate Chl-a averages for a period ranging from the early spring (when most barnacle larvae were in the water) to the summer (when barnacle size was measured after weeks of growth following spring benthic recruitment). The relationships were always positive, Chl-a explaining nearly half, or more, of the variation in barnacle size in four of the five studied years. These are remarkable results because they were based on a relatively limited number of locations (which often curtails statistical power) and point to the relevance of pelagic food supply to explain variation in intertidal barnacle size along this western ocean boundary coast.

Density of intertidal barnacles along their full elevational range of distribution conforms to the abundant-centre hypothesis.

The abundant-centre hypothesis (ACH) predicts that the density of a species should peak at its distribution centre and decrease similarly towards distribution margins. The ACH has been deduced from a theory that postulates that environmental conditions should be most favourable for a species at the centre of its distribution. This idealised density pattern, however, has been supported by limited field studies, as natural patterns are often more complex. It is thus of interest to examine under what conditions compliance with the ACH could be favoured. Such conditions could be smooth environmental gradients with limited habitat patchiness throughout the distribution range of a species. Thus, we tested the ACH by measuring the density of an intertidal barnacle (Semibalanus balanoides) across its full vertical distribution range (from low to high elevations) on a rocky shore with similar substrate properties across elevations. To do a reliable test, we surveyed eight elevation zones applying an equal sampling effort in each zone. Average barnacle density conformed to the ACH, as it peaked at the middle of the vertical distribution range of this species. The same underlying theory predicts a similar unimodal pattern for maximum body size, but this trait was decoupled from density, as maximum barnacle size increased from low to high elevations. Overall, although the ACH is not a universal predictive tool as once envisioned, it may predict some cases well, as shown by this study. Therefore, the ACH should not be discarded completely, and its domain of application should be further evaluated.

Unimodal relationship between small-scale barnacle recruitment and the density of pre-existing barnacle adults.

Recruitment is a key demographic process for population persistence. This paper focuses on barnacle (Semibalanus balanoides) recruitment. In rocky intertidal habitats from the Gulf of St. Lawrence coast of Nova Scotia (Canada), ice scour is common during the winter. At the onset of intertidal barnacle recruitment in early May (after sea ice has fully melted), mostly only adult barnacles and bare substrate are visible at high elevations in wave-exposed habitats. We conducted a multiannual study to investigate if small-scale barnacle recruitment could be predicted from the density of pre-existing adult barnacles. In a year that exhibited a wide adult density range (ca. 0-130 individuals dm-2), the relationship between adult density and recruit density (scaled to the available area for recruitment, which excluded adult barnacles) was unimodal. In years that exhibited a lower adult density range (ca. 0-40/50 individuals dm-2), the relationship between adult and recruit density was positive and resembled the lower half of the unimodal relationship. Overall, adult barnacle density was able to explain 26-40% of the observed variation in recruit density. The unimodal adult-recruit relationship is consistent with previously documented intraspecific interactions. Between low and intermediate adult densities, the positive nature of the relationship relates to the previously documented fact that settlement-seeking larvae are chemically and visually attracted to adults, which might be important for local population persistence. Between intermediate and high adult densities, where population persistence may be less compromised and the abundant adults may limit recruit growth and survival, the negative nature of the relationship suggests that adult barnacles at increasingly high densities stimulate larvae to settle elsewhere. The unimodal pattern may be especially common on shores with moderate rates of larval supply to the shore, because high rates of larval supply may swamp the coast with settlers, decoupling recruit density from local adult abundance.

A 12-year record of intertidal barnacle recruitment in Atlantic Canada (2005-2016): relationships with sea surface temperature and phytoplankton abundance.

On the Gulf of St. Lawrence coast of Nova Scotia (Canada), recruitment of the barnacle Semibalanus balanoides occurs in May and June. Every year in June between 2005 and 2016, we recorded recruit density for this barnacle at the same wave-exposed rocky intertidal location on this coast. During these 12 years, mean recruit density was lowest in 2015 (198 recruits dm-2) and highest in 2007 (969 recruits dm-2). The highest recruit density observed in a single quadrat was 1,457 recruits dm-2 (in 2011) and the lowest was 34 recruits dm-2 (in 2015). Most barnacle recruits appear during May, which suggests that most pelagic larvae (which develop over 5-6 weeks before benthic settlement) are in the water column in April. An AICc-based model selection approach identified sea surface temperature (SST) in April and the abundance of phytoplankton (food for barnacle larvae, measured as chlorophyll-a concentration -Chl-a-) in April as good explanatory variables. Together, April SST and April Chl-a explained 51% of the observed interannual variation in recruit density, with an overall positive influence. April SST was positively related to March-April air temperature (AT). April Chl-a was negatively related to the April ratio between the number of days with onshore winds (which blow from phytoplankton-limited offshore waters) and the number of days with alongshore winds (phytoplankton is more abundant on coastal waters). Therefore, this study suggests that climatic processes affecting April SST and April Chl-a indirectly influence intertidal barnacle recruitment by influencing larval performance.

Adult Prey Neutralizes Predator Nonconsumptive Limitation of Prey Recruitment.

Recent studies have shown that predator chemical cues can limit prey demographic rates such as recruitment. For instance, barnacle pelagic larvae reduce settlement where predatory dogwhelk cues are detected, thereby limiting benthic recruitment. However, adult barnacles attract conspecific larvae through chemical and visual cues, aiding larvae to find suitable habitat for development. Thus, we tested the hypothesis that the presence of adult barnacles (Semibalanus balanoides) can neutralize dogwhelk (Nucella lapillus) nonconsumptive effects on barnacle recruitment. We did a field experiment in Atlantic Canada during the 2012 and 2013 barnacle recruitment seasons (May-June). We manipulated the presence of dogwhelks (without allowing them to physically contact barnacles) and adult barnacles in cages established in rocky intertidal habitats. At the end of both recruitment seasons, we measured barnacle recruit density on tiles kept inside the cages. Without adult barnacles, the nearby presence of dogwhelks limited barnacle recruitment by 51%. However, the presence of adult barnacles increased barnacle recruitment by 44% and neutralized dogwhelk nonconsumptive effects on barnacle recruitment, as recruit density was unaffected by dogwhelk presence. For species from several invertebrate phyla, benthic adult organisms attract conspecific pelagic larvae. Thus, adult prey might commonly constitute a key factor preventing negative predator nonconsumptive effects on prey recruitment.

Predator nonconsumptive effects on prey recruitment weaken with recruit density.

We investigated the nonconsumptive effects (NCEs) of predatory dogwhelks (Nucella lapillus) on intertidal barnacle (Semibalanus balanoides) recruitment through field experiments on the Gulf of St. Lawrence coast and the Atlantic coast of Nova Scotia, Canada. We studied the recruitment seasons (May-June) of 2011 and 2013. In 2011, the Gulf coast had five times more nearshore phytoplankton (food for barnacle larvae and recruits) during the recruitment season and yielded a 58% higher barnacle recruit density than the Atlantic coast at the end of the recruitment season. In 2013, phytoplankton levels and barnacle recruit density were similar on both coasts and also lower than for the Gulf coast in 2011. Using the comparative-experimental method, the manipulation of dogwhelk presence (without allowing physical contact with prey) revealed that dogwhelk cues limited barnacle recruitment under moderate recruit densities (Atlantic 2011/2013 and Gulf 2013) but had no effect under a high recruit density (Gulf 2011). Barnacle recruits attract settling larvae through chemical cues. Thus, the highest recruit density appears to have neutralized dogwhelk effects. This study suggests that the predation risk perceived by settling larvae may decrease with increasing recruit density and that prey food supply may indirectly influence predator NCEs on prey recruitment.

Visual record of intertidal disturbance caused by drift ice in the spring on the Atlantic coast of Nova Scotia.

In the early spring of 2014, an unusually large amount of sea ice drifted from the Gulf of St. Lawrence, where it had been produced, towards the open Atlantic Ocean through the Cabot Strait, between Nova Scotia and Newfoundland, Canada. In early April, significant amounts of drift ice reached the Atlantic coast of mainland Nova Scotia. The ice floes persisted in those coastal waters for up to 16 days, depending on the location. During that time, the ice fragments caused extensive physical disturbance in rocky intertidal communities, removing high quantities of seaweeds and invertebrates. For example, at a location where the ice stayed for 9 days, the loss of macroalgal and invertebrate biomass was almost total. At a location where the ice stayed for 4 days, losses were lower, albeit still high overall. Such a magnitude of disturbance is not common on this coast, as sea ice had not reached the surveyed locations in the previous 4-5 years. We suggest that the frequency of ice scour events may help to predict intertidal community structure. This notion could be tested through multiannual surveys of ice conditions and biological communities along the Atlantic coast of Nova Scotia.

Differential recolonization of Atlantic intertidal habitats after disturbance reveals potential bottom-up community regulation.

In the spring of 2014, abundant sea ice that drifted out of the Gulf of St. Lawrence caused extensive disturbance in rocky intertidal habitats on the northern Atlantic coast of mainland Nova Scotia, Canada. To monitor recovery of intertidal communities, we surveyed two wave-exposed locations in the early summer of 2014. Barnacle recruitment and the abundance of predatory dogwhelks were low at one location (Tor Bay Provincial Park) but more than 20 times higher at the other location (Whitehead). Satellite data indicated that the abundance of coastal phytoplankton (the main food source for barnacle larvae) was consistently higher at Whitehead just before the barnacle recruitment season, when barnacle larvae were in the water column. These observations suggest bottom-up forcing of intertidal communities. The underlying mechanisms and their intensity along the NW Atlantic coast could be investigated through studies done at local and regional scales.

Variation in community structure across vertical intertidal stress gradients: how does it compare with horizontal variation at different scales?

In rocky intertidal habitats, the pronounced increase in environmental stress from low to high elevations greatly affects community structure, that is, the combined measure of species identity and their relative abundance. Recent studies have shown that ecological variation also occurs along the coastline at a variety of spatial scales. Little is known, however, on how vertical variation compares with horizontal variation measured at increasing spatial scales (in terms of sampling interval). Because broad-scale processes can generate geographical patterns in community structure, we tested the hypothesis that vertical ecological variation is higher than fine-scale horizontal variation but lower than broad-scale horizontal variation. To test this prediction, we compared the variation in community structure across intertidal elevations on rocky shores of Helgoland Island with independent estimates of horizontal variation measured at the scale of patches (quadrats separated by 10s of cm), sites (quadrats separated by a few m), and shores (quadrats separated by 100s to 1000s of m). The multivariate analyses done on community structure supported our prediction. Specifically, vertical variation was significantly higher than patch- and site-scale horizontal variation but lower than shore-scale horizontal variation. Similar patterns were found for the variation in abundance of foundation taxa such as Fucus spp. and Mastocarpus stellatus, suggesting that the effects of these canopy-forming algae, known to function as ecosystem engineers, may explain part of the observed variability in community structure. Our findings suggest that broad-scale processes affecting species performance increase ecological variability relative to the pervasive fine-scale patchiness already described for marine coasts and the well known variation caused by vertical stress gradients. Our results also indicate that experimental research aiming to understand community structure on marine shores should benefit from applying a multi-scale approach.

Positive interspecific relationship between temporal occurrence and abundance in insects.

One of the most studied macroecological patterns is the interspecific abundance-occupancy relationship, which relates species distribution and abundance across space. Interspecific relationships between temporal distribution and abundance, however, remain largely unexplored. Using data for a natural assemblage of tabanid flies measured daily during spring and summer in Nova Scotia, we found that temporal occurrence (proportion of sampling dates in which a species occurred in an experimental trap) was positively related to temporal mean abundance (number of individuals collected for a species during the study period divided by the total number of sampling dates). Moreover, two models that often describe spatial abundance-occupancy relationships well, the He-Gaston and negative binomial models, explained a high amount of the variation in our temporal data. As for the spatial abundance-occupancy relationship, the (temporal) aggregation parameter, k, emerged as an important component of the hereby named interspecific temporal abundance-occurrence relationship. This may be another case in which a macroecological pattern shows similarities across space and time, and it deserves further research because it may improve our ability to forecast colonization dynamics and biological impacts.

ROLE OF SURFACE WOUNDS AND BROWN ALGAL EPIPHYTES IN THE COLONIZATION OF ASCOPHYLLUM NODOSUM (PHAEOPHYCEAE) FRONDS BY VERTEBRATA LANOSA (RHODOPHYTA)(1).

Ascophyllum nodosum (L.) Le Jol. forms extensive beds in wave-sheltered, rocky intertidal habitats on the northwestern Atlantic coast. This fucoid seaweed is host to an obligate red algal epiphyte, Vertebrata lanosa (L.) T. A. Chr. [=Polysiphonia lanosa (L.) Tandy], and two facultative brown algal epiphytes, Elachista fucicola (Velley) Aresch. and Pylaiella littoralis (L.) Kjellm. Although V. lanosa can occur throughout most of the length of host fronds, it largely predominates in midfrond segments. The two brown algal epiphytes are restricted to distal segments. Through field experiments conducted in Nova Scotia, Canada, we tested the hypothesis that surface wounds are required for the colonization of distal segments of host fronds by V. lanosa. Distal tissues normally have a smooth surface because of their young age (A. nodosum fronds grow apically). By creating small wounds that mimicked grazing wounds distributed elsewhere on host fronds, we demonstrated that V. lanosa can colonize distal frond segments during the growth and reproductive season (summer and autumn). Approximately half of the artificial wounds were colonized by V. lanosa during this time. The experimental exclusion of both brown algal epiphytes from distal frond segments did not affect colonization by V. lanosa. Thus, we conclude that the absence of surface irregularities on distal segments of host fronds, specifically small wounds, is the main factor explaining the absence of V. lanosa there. We propose that further experimental work clarifying epiphyte distribution in host beds will enhance our ability to understand the functional role of epiphytes in intertidal ecosystems.

DISTRIBUTION OF ALGAL EPIPHYTES ACROSS ENVIRONMENTAL GRADIENTS AT DIFFERENT SCALES: INTERTIDAL ELEVATION, HOST CANOPIES, AND HOST FRONDS(1).

Understanding epiphyte distribution in coastal communities is important because these organisms affect many others directly or indirectly. Yet, their distribution has been considerably less studied than that of their hosts and other primary-space holders. Identifying major sources of variation in epiphyte abundance is thus still a need. Environmental gradients help predict species distribution and are pervasive on marine shores. In this study, we test the notion that environmental gradients across intertidal elevation, throughout host canopies, and along host fronds explain a large variation in the abundance of sympatric epiphytes. Our model system was the assemblage of Ascophyllum nodosum (L.) Le Jol. and its epiphytes Vertebrata lanosa (L.) T. A. Chr. [= Polysiphonia lanosa (L.) Tandy], Elachista fucicola (Velley) Aresch., and Pylaiella littoralis (L.) Kjellm. On the coast of Nova Scotia (Canada), we found evidence of a spatial segregation among these species at almost all scales. While the red epiphyte V. lanosa was more common at high- and midintertidal elevations (peaking at midelevations) and on middle segments of host fronds, the brown epiphytes E. fucicola and P. littoralis were more common at low elevations and restricted to distal segments of host fronds. Canopy habitat affected abundance only for V. lanosa, which was more common within the host canopy than on its periphery at midelevations. Since the studied gradients are related to predictable changes in abiotic factors, the identification of likely causes behind the observed patterns was facilitated. Our study ends by proposing abiotic and biotic factors that deserve priority in the experimental testing of the forces structuring this assemblage.