The role of prey size and abundance in the geographical distribution of spider sociality

1. Social species in the spider genus Anelosimus predominate in lowland tropical rainforests, while congeneric subsocial species occur at higher elevations or higher latitudes. 2. We conducted a comparative study to determine whether differences in total biomass, insect size or both have been responsible for this pattern. 3. We found that larger average insect size, rather than greater overall biomass per se, is a key characteristic of lowland tropical habitats correlating with greater sociality. 4. Social species occupied environments with insects several times larger than the spiders, while subsocial species nearing dispersal occupied environments with smaller insects in either high or low overall biomass. 5. Similarly, in subsocial spider colonies, individuals lived communally at a time when they were younger and therefore smaller than the average insect landing on their webs. 6. We thus suggest that the availability of large insects may be a critical factor restricting social species to their lowland tropical habitats.     

Altitudinal patterns of spider sociality and the biology of a new midelevation social Anelosimus species in Ecuador

To the extent that geography correlates with particular environmental parameters, the geographical distribution of phylogenetically related social and nonsocial organisms should shed light on the conditions that lead to sociality versus nonsociality. Social spiders are notorious for being concentrated in tropical regions of the world, occupying a set of habitats more restricted than those available to the phylogenetic lineages in which they occur. Here we document a parallel pattern involving elevation in the spider genus Anelosimus in America and describe the biology of a newly discovered social species found at what appears to be the altitudinal edge of sociality in the genus. We show that this is a cooperative permanent-social species with highly female-biased sex ratios but colonies that are one to two orders of magnitude smaller than those of a low-elevation congener of similar body size. We suggest that the absence of subsocial Anelosimus species in the lowland rain forest may be due to an increased probability of maternal death in this habitat due to greater predation and/or precipitation, while absence of a sufficient supply of large insects at high elevations or latitudes may restrict social species to low- to midelevation tropical moist forests. We refer to these as the "maternal survival" and "prey size" hypotheses, respectively, and suggest that both in combination may explain the geographical distribution of sociality in the genus.     

Revision and phylogenetic analysis of American ethicus and rupununi groups of Anelosimus (Araneae, Theridiidae)

Two species groups of the social spider genus Anelosimus are revised. The ethicus group contains six species found in South America, in an area ranging from the Guianas to southern Brazil and Argentina. Of these, A. rabus Levi, 1963, A. ethicus ( Keyserling, 1884 ), and A. nigrescens ( Keyserling, 1884 ) are redescribed, while A. nigrescens is removed from synonymy with A. ethicus . Three new species are described: A. misiones sp. nov., A. sumisolena sp. nov. and A. inhandava sp. nov. Anelosimus ethicus is reportedly either subsocial or solitary, while the behaviour of the other species in the group is unknown. The rupununi group contains two quasisocial species, A. rupununi Levi, 1956 and A. lorenzo Fowler & Levi, 1979, from the Caribbean and tropical South America. Both are redescribed here. A parsimony analysis of morphological characters provides support for the monophyly of both groups. In the phylogeny, subsociality optimizes to the base of Anelosimus , indicating that the common ancestor of the ethicus group was subsocial. Its members can thus be predicted to be subsocial, or secondarily solitary. Quasisociality arose de novo in the rupununi group, representing one of 6−7 independent origins in theridiids. Study of the biology of Anelosimus is important to advance our understanding of the evolution of sociality in spiders.     

Group living and inbreeding depression in a subsocial spider

Social spiders are unusual among social organisms in being highly inbred-males and females mature within their natal nest and mate with each other to produce successive generations. Several lines of evidence suggest that in spiders inbred social species originated from outbred subsocial ancestors, a transition expected to have been hindered by inbreeding depression. As a window into this transition, we examined the fitness consequences of artificially imposed inbreeding in the naturally outbred subsocial spider Anelosimus cf. jucundus. Subsocial spiders alternate periods of solitary and social living and are thought to resemble the ancestral system from which the inbred social species originated. We found that inbreeding depression in this subsocial spider only becomes evident in spiders raised individually following the end of their social phase and that ecological and demographic factors such as eclosion date, number of siblings in the group and mother's persistence are more powerful determinants of fitness during the social phase. A potential explanation for this pattern is that maternal care and group living provide a buffer against inbreeding depression, a possibility that may help explain the repeated origin of inbred social systems in spiders and shed light on the origin of other systems involving regular inbreeding.     

Subsocial spiders in space and time: A fine scale approach to the dynamics of dispersal

Less than 0.2% of all spider species live in close associations with conspecifics. Among these, subsocial spiders show characteristics of both solitary spiders (e.g., individuals disperse for breeding) and social spiders (e.g., prolonged cooperative behaviours at least prior to independent reproduction). Dispersing individuals build small webs, usually with one inhabitant, whereas colonies are large webs with plant debris and harbouring multiple females. We studied the spatiotemporal dynamics of dispersal in the subsocial spider Anelosimus baeza. We followed the occupancy of all colonies and dispersal webs over the breeding season by mapping the number and sex of spiders with respect to their location in three dimensions. We studied the settlement patterns of new webs and fluctuation in web occupancy through movement between occupied and abandoned webs of colonies and dispersal webs. The occupancy of webs was highly dynamic with changes occurring at small time scales. The similarity in the patterns of web occupancy by females among dispersal webs was partially explained by their spatial and their temporal proximity. Our results suggest that dispersal webs may be used by spiders as a temporary refuge by both sexes during the breeding season. Patterns described here suggest new approaches to dispersal studies in group living spiders.     

Iterative evolution of increased behavioral variation characterizes the transition to sociality in spiders and proves advantageous

Abstract The evolution of group living is regarded as a major evolutionary transition and is commonly met with correlated shifts in ancillary characters. We tested for associations between social tendency and a myriad of abiotic variables (e.g., temperature and precipitation) and behavioral traits (e.g., boldness, activity level, and aggression) in a clade of spiders that exhibit highly variable social structures (genus Anelosimus). We found that, relative to their subsocial relatives, social species tended to exhibit reduced aggressiveness toward prey, increased fearfulness toward predators, and reduced activity levels, and they tended to occur in warm, wet habitats with low average wind velocities. Within-species variation in aggressiveness and boldness was also positively associated with sociality. We then assessed the functional consequences of within-species trait variation on reconstituted colonies of four test species (Anelosimus eximius, Anelosimus rupununi, Anelosimus guacamayos, and Anelosimus oritoyacu). We used colonies consisting of known ratios of docile versus aggressive individuals and group foraging success as a measure of colony performance. In all four test species, we found that groups composed of a mixture of docile and aggressive individuals outperformed monotypic groups. Mixed groups were more effective at subduing medium and large prey, and mixed groups collectively gained more mass during shared feeding events. Our results suggest that the iterative evolution of depressed aggressiveness and increased within-species behavioral variation in social spiders is advantageous and could be an adaptation to group living that is analogous to the formation of morphological castes within the social insects.     

Shifting continents, not behaviours: independent colonization of solitary and subsocial Anelosimus spider lineages on Madagascar (Araneae, Theridiidae)

Agnarsson, I., Kuntner, M., Coddington, J. A. & Blackledge, T. A. (2009). Shifting continents, not behaviours: independent colonization of solitary and subsocial Anelosimus spider lineages on Madagascar (Araneae, Theridiidae). —Zoologica Scripta, 39, 75–87. Madagascar is a biodiversity hotspot, thought to be colonized mostly via Cenozoic dispersal from Africa, followed by endemic radiation of multiple lineages. Anelosimus spiders are diverse in Madagascar, and, like their congeners in the Americas, are most diverse in wet montane forests. Most Anelosimus species are social in that they cooperate in web building and prey capture either during a part of their life cycles (subsocial), including hitherto studied Malagasy species, or permanently (quasisocial). One Central American coastal species, Anelosimus pacificus, has secondarily switched to solitary living, and available evidence suggests that its closest relatives from S. America and Europe are likely also solitary. Here, we show that the only known coastal Anelosimus species in Madagascar and Comoros –Anelosimus decaryi and Anelosimus amelie sp. n. – are also solitary. Using a phylogenetic approach, we test two competing hypotheses: (i) that Malagasy Anelosimus are monophyletic and thus represent a second example of reversal to solitary living in a littoral habitat or (ii) that solitary and subsocial lineages independently colonized Madagascar. We find that solitary Malagasy Anelosimus are closely related to their solitary counterparts from Europe and the Americas, while subsocial Malagasy species nest sister to Anelosimus nelsoni from S. Africa. This finding suggests that (i) the two Anelosimus lineages colonized Madagascar independently and (ii) a reversal to solitary behaviour has occurred only once in Anelosimus. Thus, solitary littoral Malagasy species did not descend from Malagasy mountains, but arrived from much further afar. African and possibly American origin of the two lineages is implied by our findings. To restore natural classification of Anelosimus, Seycellocesa Koçak & Kemal is synonymized with it.     

Smaller colonies and more solitary living mark higher elevation populations of a social spider

1. There appears to be a pattern of decreasing sociality with increasing elevation across social spider species in the genus Anelosimus at tropical latitudes. Our data suggest that this pattern holds within a single species, Anelosimus eximius, on a smaller altitudinal gradient. 2. In comparing colony size at six different altitudes in north-eastern Ecuador, we find that the lowland A. eximius populations tend to have larger colonies and few solitary females. At higher elevations, many of the colonies are small and the proportion of solitary females is greater. 3. Contrary to expectation, we also found no difference in spider density between the upper elevation and lowland populations. This result may be partly due to the fact that upper elevation populations occur only at the forest edge (as opposed to both edge and interior) where populations at all elevations appear more robust.     

Utilization of Sugarcane Habitat by Feral Pig (Sus scrofa) in Northern Tropical Queensland: Evidence from the Stable Isotope Composition of Hair

Feral pigs (Sus scrofa) are an invasive species that disrupt ecosystem functioning throughout their introduced range. In tropical environments, feral pigs are associated with predation and displacement of endangered species, modification of habitat, and act as a vector for the spread of exotic vegetation and disease. Across many parts of their introduced range, the diet of feral pigs is poorly known. Although the remote location and difficult terrain of far north Queensland makes observing feral pig behavior difficult, feral pigs are perceived to seek refuge in World Heritage tropical rainforests and seasonally 'crop raid' into lowland sugarcane crops. Thus, identifying how feral pigs are using different components of the landscape is important to the design of management strategies. We used the stable isotope composition of captured feral pigs to determine the extent of rainforest and sugarcane habitat usage. Recently grown hair (basal hair) from feral pigs captured in remote rainforest indicated pigs met their dietary needs solely within this habitat. Stable carbon and nitrogen isotope values of basal hair from feral pigs captured near sugarcane plantations were more variable, with some individuals estimated to consume over 85% of their diet within a sugarcane habitat, while a few consumed as much as 90% of their diet from adjacent forested environments. We estimated whether feral pigs switch habitats by sequentially sampling δ(13)C and δ(15)N values of long tail hair from a subset of seven captured animals, and demonstrate that four of these individuals moved between habitats. Our results indicate that feral pigs utilize both sugarcane and forest habitats, and can switch between these resources.     

Sensitivity of tropical forests to climate change in the humid tropics of north Queensland

An analysis using an artificial neural network model suggests that the tropical forests of north Queensland are highly sensitive to climate change within the range that is likely to occur in the next 50–100 years. The distribution and extent of environments suitable for 15 structural forest types were estimated, using the model, in 10 climate scenarios that include warming up to 1°C and altered precipitation from –10% to +20%. Large changes in the distribution of forest environments are predicted with even minor climate change. Increased precipitation favours some rainforest types, whereas decreased rainfall increases the area suitable for forests dominated by sclerophyllous genera such as Eucalyptus and Allocasuarina. Rainforest environments respond differentially to increased temperature. The area of lowland mesophyll vine forest environments increases with warming, whereas upland complex notophyll vine forest environments respond either positively or negatively to temperature, depending on precipitation. Highland rainforest environments (simple notophyll and simple microphyll vine fern forests and thickets), the habitat for many of the region’s endemic vertebrates, decrease by 50% with only a 1°C warming. Estimates of the stress to present forests resulting from spatial shifts of forest environments (assuming no change in the present forest distributions) indicate that several forest types would be highly stressed by a 1°C warming and most are sensitive to any change in rainfall. Most forests will experience climates in the near future that are more appropriate to some other structural forest type. Thus, the propensity for ecological change in the region is high and, in the long term, significant shifts in the extent and spatial distribution of forests are likely. A detailed spatial analysis of the sensitivity to climate change indicates that the strongest effects of climate change will be experienced at boundaries between forest classes and in ecotonal communities between rainforest and open woodland.     

Kin recognition and cannibalism in a subsocial spider

Evolution of cooperation and group living in spiders from subsocial family groups may be constrained by their cannibalistic nature. A tendency to avoid cannibalizing kin may facilitate tolerance among spiders and implies the ability to identify relatives. We investigated whether the subsocial spider Stegodyphus lineatus discriminates kin by recording cannibalism among juveniles in experiments during which amount of food and size difference among spiders in groups were varied. We hypothesized that family groups should be less cannibalistic than groups of mixed‐parental origin. Further, we tested whether food‐stress would influence cannibalism rates differently in kin and nonkin groups and the effect of relatedness on cannibalism within groups of spiders of variable size compared with those of homogenous size. In groups of six spiders, more spiders were cannibalized in nonsib groups than in sib groups under low food conditions. A tendency for nonkin biased cannibalism in starved spider pairs supported that kin recognition in S. lineatus is expressed when food is limited. Size variance of individuals within well‐fed groups of siblings and unrelated spiders had no influence on cannibalism rates. Apparently, both hunger and high density are important promoters of cannibalism. In addition to inclusive fitness benefits, we suggest that an ability to avoid cannibalizing kin will favour the evolution of cooperation and group living in phylogenetically pre‐adapted solitary species.     

Natal Dispersal Patterns of a Subsocial Spider Anelosimus cf. jucundus (Theridiidae)

Species that alternate periods of solitary and social living may provide clues to the conditions that favor sociality. Social spiders probably originated from subsocial‐like ancestors, species in which siblings remain together for part of their life cycle but disperse prior to mating. Exploring the factors that lead to dispersal in subsocial species, but allow the development of large multigenerational colonies in social species, may provide insight into this transition. We studied the natal dispersal patterns of a subsocial spider, Anelosimus cf. jucundus, in Southeastern Arizona. In this population, spiders disperse from their natal nests in their penultimate and antepenultimate instars over a 3‐mo period. We tracked the natal dispersal of marked spiders at sites with clustered vs. isolated nests. We found that most spiders initially dispersed less than 5 m from their natal nests. Males and females, and spiders in patches with different densities of nests, dispersed similar distances. The fact that both sexes in a group dispersed, the lack of a sex difference in dispersal distance, and the relatively short distances dispersed are consistent with the hypothesis that natal dispersal results from resource competition within the natal nest, rather than inbreeding avoidance in competition for mates. Additionally, an increase in the average distance dispersed with time and with the number of spiders leaving a nest suggests that competition for nest sites in the vicinity of the natal nest may affect dispersal distances. The similar distances dispersed in patches with isolated vs. clustered nests, in contrast, suggest that competition among dispersers from different nests may not affect dispersal distances.     

Immigrants and refugees: the importance of dispersal in mediating biotic attrition under climate change

Montane tropical rainforests are critically important areas for global bird diversity, but are projected to be highly vulnerable to contemporary climate change. Upslope shifts of lowland species may partially offset declines in upland species but also result in a process of lowland biotic attrition. This latter process is contingent on the absence of species adapted to novel warm climates, and isolation from pools of potential colonizers. In the Australian Wet Tropics, species distribution modelling has forecast critical declines in suitable environmental area for upland endemic birds, raising the question of the future role of both natural and assisted dispersal in species survival, but information is lacking for important neighbouring rainforest regions. Here we use expanded geographic coverage of data to model the realized distributions of 120 bird species found in north‐eastern Australian rainforest, including species from potential source locations in the north and recipient locations in the south. We reaffirm previous conclusions as to the high vulnerability of this fauna to global warming, and extend the list of species whose suitable environmental area is projected to decrease. However, we find that expansion of suitable area for some species currently restricted to northern rainforests has the potential to offset biotic attrition in lowland forest of the Australian Wet Tropics. By examining contrasting dispersal scenarios, we show that responses to climate change in this region may critically depend on dispersal limitation, as climate change shifts the suitable environmental envelopes of many species south into currently unsuitable habitats. For lowland and northern species, future change in vegetation connectivity across contemporary habitat barriers is likely to be an important mediator of climate change impacts. In contrast, upland species are projected to become increasingly isolated and restricted. Their survival is likely to be more dependent on the viability of assisted migration, and the emergence and persistence of suitable environments at recipient locations.     

Advantages of social foraging in crab spiders: Groups capture more and larger prey despite the absence of a web

Among group‐living spiders, subsocial representatives in the family of crab spiders (Thomisidae) are a special case, as they build protective communal leaf nests instead of extensive communal capture webs. It could thus be inferred that antipredator benefits (e.g., enhanced protection in larger nests) rather than foraging‐related advantages (e.g., capture of more and larger prey) promote sociality in this family. Nonetheless, subsocial crab spiders do share prey, and if this behaviour does not reflect mere food scramble but has a cooperative character, crab spiders may offer insights into the evolution of social foraging applicable to many other cooperative predators that hunt without traps. Here, we performed a comparative laboratory feeding experiment on three of the four subsocial crab spider species—Australomisidia ergandros, Australomisidia socialis and Xysticus bimaculatus—to determine if crab spiders derive advantages from foraging in groups. In particular, we tested artificially composed groups of five sibling spiderlings vs. single siblings in terms of prey capture success and prey size preference. Across species, groups had higher prey capture success (measured in terms of capture rates and capture latency) and were more likely to attack large, sharable prey—dynamics leading to reduced food competition among group members in favour of living and foraging in groups. Within groups, we further compared prey extraction efficiency among the three applied social foraging tactics: producing, scrounging and feeding alone. In A. ergandros, individuals were exceptionally efficient when using the non‐cooperative scrounger tactic, which entails feeding on the prey provided by others. Thus, our multispecies comparison confirms foraging advantages in maintaining a cooperative lifestyle for crab spiders, but also demonstrates the relevance of research into exploitation of cooperative foraging in this family.     

The evolution of social inbreeding mating systems in spiders: limited male mating dispersal and lack of pre‐copulatory inbreeding avoidance in a subsocial predecessor

Cooperation and group living are extremely rare in spiders and only few species are known to be permanently social. Inbreeding is a key characteristic of social spiders, resulting in high degrees of within‐colony relatedness that may foster kin‐selected benefits of cooperation. Accordingly, philopatry and regular inbreeding are suggested to play a major role in the repeated independent origins of sociality in spiders. We conducted field observations and laboratory experiments to investigate the mating system of the subsocial spider Stegodyphus tentoriicola. The species is suggested to resemble the ‘missing link’ in the transition from subsociality to permanent sociality in Stegodyphus spiders because its social period is prolonged in comparison to other subsocial species. Individuals in our two study populations were spatially clustered around maternal nests, indicating that clusters consist of family groups as found in the subsocial congener Stegodyphus lineatus. Male mating dispersal was limited and we found no obvious pre‐copulatory inbreeding avoidance, suggesting a high likelihood of mating with close kin. Rates of polygamy were low, a pattern ensuring high relatedness within broods. In combination with ecological constraints, such as high costs of dispersal, our findings are consistent with the hypothesis that the extended social period in S. tentoriicola is accompanied with adaptations that facilitate the transition towards permanent sociality. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 851–859.     

Social behaviour in a web-building lynx spider, Tapinillus sp. (Araneae: Oxyopidae)

Social behaviour involving cooperative prey capture and communal feeding is reported for the first time in the spider family Oxyopidae (lynx spiders), in a web-building species of the genus Tapinillus. This social spider inhabits communal webs that may contain several dozen individuals, including adults of both sexes and juveniles of different cohorts. Its colonies occur in clusters and appear to be long-lived, much like those of non-territorial permanently social species such as Anelosimus eximius (Theridiidae) or Agelena consociata (Agelenidae). However, unlike colonies of these other cooperative spiders, the colonies of the social Tapinillus do not have highly female-biased sex ratios. The possible explanations for this difference are discussed.     

Effects of rainforest fragmentation and shade-coffee plantations on spider communities in the Western Ghats,India

Studies on the effects of tropical rainforest fragmentation and disturbance have often focussed on plants and vertebrates such as birds and mammals and seldom on invertebrates, despite the latter being among the most biologically diverse groups in these ecosystems. Spiders are one such group of invertebrate predators that are known to be sensitive indicators of environmental change in tropical ecosystems. The present study assesses the spider community structure and responses to rainforest fragmentation and degradation and conversion to shade-coffee plantations in the Anamalai hills, southern Western Ghats, India. Ten rainforest fragments ranging in size from 11 ha to 2,600 ha under varying levels of degradation within the Indira Gandhi Wildlife Sanctuary and private lands of the Valparai plateau, and two shade-coffee plantation sites were sampled for spiders using visual searches along time-constrained belt transects between January and May 2005. Within a total sampled area of 5.76 ha, 4,565 individual spiders (4,300 detections) belonging to 156 morphospecies within 21 families and 8 functional groups were recorded. The estimated total number of understorey spider species in the study area was 192 (±5.15 SD) species, representing around 13% of the total number of spider species so far described from India. Overall spider density, species richness, and species density showed no trend in relation to fragment area across all sites. Specific comparisons among undisturbed sites indicated however that high altitude sites had fewer species than mid-altitude sites and fragments had fewer species than relatively larger continuous forest sites. In contrast to the lack of trend in overall species richness and abundance, species composition changed substantially in relation to habitat alteration and altitude. Cluster analysis of Bray-Curtis similarities among sites in spider species composition revealed four distinct clusters: high altitude undisturbed sites, mid-altitude disturbed sites with an undisturbed mid altitude site, mid-altitude highly disturbed sites with a disturbed site, and shade-coffee plantation sites. Spider species, such as Psechrus torvus and Tylorida culta, that contributed significantly to the dissimilarity between undisturbed and disturbed rainforest sites, and rainforest and shade-coffee sites were identified that serve as useful indicators of habitat alteration.     

The transition to social inbred mating systems in spiders: role of inbreeding tolerance in a subsocial predecessor

The social spiders are unusual among cooperatively breeding animals in being highly inbred. In contrast, most other social organisms are outbred owing to inbreeding avoidance mechanisms. The social spiders appear to originate from solitary subsocial ancestors, implying a transition from outbreeding to inbreeding mating systems. Such a transition may be constrained by inbreeding avoidance tactics or fitness loss due to inbreeding depression. We examined whether the mating system of a subsocial spider, in a genus with three social congeners, is likely to facilitate or hinder the transition to inbreeding social systems. Populations of subsocial Stegodyphus lineatus are substructured and spiders occur in patches, which may consist of kin groups. We investigated whether male mating dispersal prevents matings within kin groups in natural populations. Approximately half of the marked males that were recovered made short moves (< 5m) and mated within their natal patch. This potential for inbreeding was counterbalanced by a relatively high proportion of immigrant males. In mating experiments, we tested whether inbreeding actually results in lower offspring fitness. Two levels of inbreeding were tested: full sibling versus non-sib matings and matings of individuals within and between naturally occurring patches of spiders. Neither full siblings nor patch mates were discriminated against as mates. Sibling matings had no effect on direct fitness traits such as fecundity, hatching success, time to hatching and survival of the offspring, but negatively affected offspring growth rates and adult body size of both males and females. Neither direct nor indirect fitness measures differed significantly between within patch and between-patch pairs. We tested the relatedness between patch mates and nonpatch mates using DNA fingerprinting (TE-AFLP). Kinship explained 30% of the genetic variation among patches, confirming that patches are often composed of kin. Overall, we found limited male dispersal, lack of kin discrimination, and tolerance to low levels of inbreeding. These results suggest a history of inbreeding which may reduce the frequency of deleterious recessive alleles in the population and promote the evolution of inbreeding tolerance. It is likely that the lack of inbreeding avoidance in subsocial predecessors has facilitated the transition to regular inbreeding social systems.     

There's no place like home: seedling mortality contributes to the habitat specialisation of tree species across Amazonia

Understanding the mechanisms generating species distributions remains a challenge, especially in hyperdiverse tropical forests. We evaluated the role of rainfall variation, soil gradients and herbivory on seedling mortality, and how variation in seedling performance along these gradients contributes to habitat specialisation. In a 4‐year experiment, replicated at the two extremes of the Amazon basin, we reciprocally transplanted 4638 tree seedlings of 41 habitat‐specialist species from seven phylogenetic lineages among the three most important forest habitats of lowland Amazonia. Rainfall variation, flooding and soil gradients strongly influenced seedling mortality, whereas herbivory had negligible impact. Seedling mortality varied strongly among habitats, consistent with predictions for habitat specialists in most lineages. This suggests that seedling performance is a primary determinant of the habitat associations of adult trees across Amazonia. It further suggests that tree diversity, currently mostly harboured in terra firme forests, may be strongly impacted by the predicted climate changes in Amazonia.