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Ecosphere Volume 10 ,Issue 3 ,2019-03-18
Birds of a feather flock together: Functionally similar vertebrates positively co‐occur in Guianan forests
Thomas Denis 1 , 2 Cécile Richard‐Hansen 1 Olivier Brunaux 3 Stéphane Guitet 3 , 4 Bruno Hérault 5 , 6
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Received 2018-09-11, accepted for publication 2018-10-19, Published 2018-10-19

Abstract Medium‐ and large‐sized vertebrates play a key role in shaping overall forest functioning. Despite this, vertebrate interactions, from competition to mutualism, remain poorly studied, even though these interactions should be taken into account in our conservation and management strategies. Thus, we tackled the question of vertebrate co‐occurrence in tropical rainforests: Are (negative or positive) co‐occurrences dependent on forest structure and composition? and Are these co‐occurrences linked to functional species similarity? We recorded the occurrence of 21 medium‐ and large‐sized vertebrates in 19 French Guianan locations in which a large set of forest structure and composition descriptors were collected. We used a probabilistic model to look for co‐occurrences at different spatial scales, and species pairwise co‐occurrences were then compared to those generated solely on the basis of forest structure and composition. We then quantified the co‐occurrence strength between pairwise species dyads and determined whether they relied on species functional similarity, controlling for the environmental effects. We found that positive co‐occurrences vastly outnumbered negative co‐occurrences, were only partly shaped by the local environment, and were closely linked to species functional similarity. Thus, groups of species sharing similar functional traits are more prone to co‐occur, highlighting the key role of functional redundancy in structuring species assemblages. We discuss how positive interactions could generate the predominance of positive co‐occurrences in oligotrophic terra firme (unflooded) forests when resources are scarce and dispersed in dry season. Finally, we identified functional groups based on co‐occurrence strength and suggested that frugivory/granivory and body size are of primary importance in species interactions in Neotropical vertebrate communities.


terra firme;mutualism;mixed‐species associations;mammals;information exchange;Guiana Shield;birds;activity matching


© 2019 The Ecological Society of America


Study area in French Guiana, northern South America, including 19 survey sites. The left part of the figure illustrates the sampling design: Four line transects were used to sample the diurnal medium‐ and large‐sized vertebrates (100‐m transect unit) and environmental conditions (i.e., forest structure and forest composition measured in each plot of 100 m × 20 m). Data were aggregated to calculate species occurrence and environmental conditions for different spatial scales from 200 to 3000 m.

Comparison of the percentage of significant positive (red) and negative (blue) co‐occurrence types between pairwise species for different spatial scales. Dashed line is the percentage of significant pairwise species co‐occurrences at P = 0.05 (significant level). Lower and upper borders of envelopes are the 0.01 and 0.10 significant levels, respectively.

Comparison of the observed number of negative (A) and positive (B) co‐occurrences between pairwise species and the expected number (5th–95th quantile) under environmentally constrained models from forest structure (solid envelopes) or forest composition (dashed envelopes) environmental covariates.

Influence of species functional dissimilarity on the co‐occurrence strength of unconstrained models and environmentally constrained models generated by forest structure. The weighted Spearman's r of each observed co‐occurrence type is compared with those of the 5th–95th quantile ranges of distributions of the environmentally constrained models generated by forest structure (central panel). See similar results for forest composition (Appendix S1: Fig. S3). Top (negative co‐occurrence) and bottom (positive co‐occurrence) panels illustrate the observed relationship, from which the Spearman's r was calculated between the species dissimilarity and the co‐occurrence strength for the 200‐, 400‐, 900‐, and 3000‐m plots. The co‐occurrence strength was calculated as the difference between the observed and expected number of plots that the two species i and j co‐occur (Jijobs‐Jijexp; Veech 2014).

Significance of univariate relationships between positive species co‐occurrence strength and trait similarity. Levels of significance are shown for the environmentally constrained model at different spatial scales and are given for the forest structure and forest composition. Circle color corresponds to the component of the environmental conditions (forest structure and composition), and the circle size to the significance level of the relationship.

(A) Cluster dendrogram of overall vertebrate assemblage based on Jaccard distance of species presence/absence. (B) Schematic illustration of presumed interspecific interactions (commensalism and mutualism). Positive co‐occurrences between species pairwise are represented by red lines. We support that terrestrial species benefit by commensalism from feeding, foraging, and other behaviors of arboreal species. Solid and dashed lines represent positive co‐occurrences within species groups and between species groups, respectively. Find after full common names sorted by initial letters and corresponding scientific names: Amazonian brown brocket deer, Mazama nemorivaga (F. Cuvier, 1817); Black curassow, Crax alector (Linnaeus, 1766); Black spider monkey, Ateles paniscus (Linnaeus, 1758); Collared peccary, Pecari tajacu (Linnaeus, 1758); Common squirrel monkey, Saimiri sciureus (Linnaeus, 1758); Guianan brown capuchin, Sapajus [Cebus] apella (Linnaeus, 1758); Golden‐handed tamarin, Saguinus midas (Linnaeus, 1758); Great Tinamou, Tinamus major (Gmelin, 1789); Guianan weeper capuchin, Cebus olivaceus (Schomburgk, 1848); Grey‐winged trumpeter, Psophia crepitans (Linnaeus, 1758); Lowland tapir, Tapirus terrestris (Linnaeus, 1758); Little tinamous, Crypturellus spp.; Marail guan, Penelope marail (Müller, 1776); Marbled wood‐quail, Odontophorus gujanensis (Gmelin, 1789); Red acouchi, Myoprocta acouchy (Erxleben, 1777); Red brocket, Mazama americana (Erxleben, 1777); Red howler monkey, Alouatta macconnelli (Linnaeus, 1766); Red‐rumped agouti, Dasyprocta leporina (Linnaeus, 1758); Tayra, Eira barbara (Linnaeus, 1758); White‐faced saki, Pithecia pithecia (Linnaeus, 1766); Yellow‐footed tortoise, Geochelone denticulata (Linnaeus, 1766).

Table 1


Thomas Denis.Office National de la Chasse et de la Faune Sauvage, UMR EcoFoG (AgroParisTech, Cirad, CNRS, INRA, Université des Antilles, Université de Guyane), 97310, Kourou, French Guiana, France;Université de Guyane, UMR EcoFoG, 97310, Kourou, French Guiana, France.thomas.denis@ecofog.gf


Thomas Denis,Cécile Richard‐Hansen,Olivier Brunaux,Stéphane Guitet,Bruno Hérault. Birds of a feather flock together: Functionally similar vertebrates positively co‐occur in Guianan forests. Ecosphere ,Vol.10, Issue 3(2019)



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