DOI: 10.5061/dryad.q83bk3jq2
title: Data from: Repeated, irreversible evolution of ant-following behavior across Neotropical avian families

description: Ecological specialization is a result of the interplay between ecological
 and evolutionary processes. One iconic ecological specialization of the
 Neotropics involves birds that follow army ant swarms in feeding groups.
 Prior work has focused on a single avian family, the Neotropical antbirds
 (Thamnophilidae), but over a century of fieldwork has now revealed that
 ant-following occurs in hundreds of distantly related birds. To understand
 the relative contributions of shared ancestry and ecological
 specialization in the evolution of ant-following, we compiled a database
 of all Neotropical ant-following birds (n = 472 species) and their degree
 of specialization on army ants, and tested if: 1) ant-following becomes
 increasingly specialized through evolutionary time, and 2)
 ecomorphological functional traits predict ant-following behavior.
 Ancestral state reconstruction revealed that specialized ant-following
 evolved independently in eight clades and four families of Neotropical
 birds (Antbirds: Thamnophilidae, Ovenbirds: Furnariidae, Tanagers:
 Thraupidae, and Cuckoos: Cuculidae). Ant-following behavior was highly
 conserved phylogenetically (Pagel’s λ = 0.97), and specialized clades
 evolved from less specialized ancestors, with few evolutionary reversals.
 In contrast, ecomorphological traits poorly predicted the level of
 ant-following specialization across species. Our results suggest
 increasing specialization on army ants is governed by niche conservatism,
 not ecological specialization.
description: # Data from: Repeated, irreversible evolution of ant-following behavior
 across Neotropical avian families #### Data and Code File Overview This
 data repository consists of 3 data files, 1 code script, and this README
 document, with the following data and code filenames and variables. **Data
 Files** 1. "FollowersOnly.csv" is a file that contains all study
 species' Latin names (column 1, 'latin_name') and their
 respective ant-following level (column 2, 'following_order') as
 an ordinal value (2-5). 2. "FollowingPhylogeny.nex" is a set of
 1000 phylogenetic trees, of object type multiphylo, of the 465 study
 species as downloaded from Vertlife.org (Jetz et al., 2012). 3.
 "Follower_Traits.csv" is the functional trait database. The
 first column ('common_name') in this database is the species
 common (English name). The second column ('latin_name') in this
 database is the latin name of the species formatted
 "Genus_species". The third column ('latin name') is
 also the latin name of the species, but it is formatted "Genus
 species". The fourth column ('elevation_min') is the
 minimum elevation of the species in meters (m) and the fifth column
 ('elevation_max') is the maximum elevation of the species in
 meters (m). The sixth column ('following_ordinal') is the
 ordinal value of ant-following, on a scale of 2 (occasional)- 6(obligate).
 The seventh column ('average*mass') is the average mass of the
 species in grams. The eight column ('Beak.Length*Culmen') is the
 beak length to the nearest 0.1mm. The ninth column
 ('Tarsus.Length') is the tarsus length to the nearest 0.1mm. The
 tenth column ('ForStrat-watbelowsurf') is the proportion of time
 foraging below the water surface. The eleventh column
 ('ForStrat-wataroundsurf') is the proportion of time foraging at
 the water surface. The twelfth column ('ForStrat-ground) is the
 proportion of time foraging on the ground. The thirteenth column
 ('ForStrat-understory') is the proportion of time foraging in
 the understory. The fourteenth column ('ForStrat-midhigh') is
 the proportion of time foraging in the midstory. The fifteenth column
 ('ForStrat-canopy') is the proportion of time foraging in the
 canopy. The sixteenth column ('ForStrat-aerial') is the
 proportion of time foraging in the air. The seventeenth column
 ('For_strat_weight') is a weighted average of the foraging
 columns (columns 10-16) to which gives the average foraging stratum of a
 species. **Code and Workflow** 1.
 "EvolutionAndTraitsAntFollowing.R"- the single R script to code
 for all of the analyses. This single R script can be run through in order.
 Generally, it is broken down into two sections. The first section is the
 analysis for the Pagel's Lambda followed by the ancestral state
 reconstructions. The analysis for the ancestral state reconstructions runs
 all of the tested models across all of the 1000 phylogenetic trees. The
 BIC and LogLikelihood values for each of these are saved and the mean and
 standard error for all BIC and LogLikelihood are calculated. Additionally
 the mean and standard error for the two different transition rates for the
 top model, ASY15 are calculated. After all the ancestral state
 reconstruction models are run, there is the code for picking the top
 model. After this, there is the code to make the reconstruction figure.
 After this, there is the code to pull out the estimated ancestral states
 of the common ancestor and basal common ancestor of the clades of interest
 in the top model-tree combination. Next, is the second section of the
 analysis. This is where the analyses to see how traits may impact
 following level. First is the code to recreate the random forest analysis
 to evaluate the classification accuracy of species army ant following
 specialization levels based on ecomorphological traits. And, lastly is the
 code to generate figure showing the position of species in each
 specialization level based on its traits. ## Software **Software** * R
 version 4.1.1 (2021-08-10) run on R Studio (2022.07.2+576). **Packages** *
 phytools (0.7-90) * picante (1.8.2) * ape (5.5) * randomForest (4.7-1.1) *
 vegan (2.5-7) * tidyverse (1.3.1) * caret (6.0-90) * lubridate * magrittr
 * ggplot2 * ggthemes * ape * phylolm * phytools * modelr * ggdist *
 viridis * logihist * gridExtra * ape * tictoc * GGally * abind * EnvStats
 * psych * rphylopic * png * scico * psych **Data Derived From Other
 Sources** The data used for this manuscript were derived from other
 sources. Below is a list of data types used in this manuscript and the
 sources from which the data were collected. Phylogenetic Data *
 Vertlife.org * Jetz, W., G. H. Thoma, J. B. Joy, K. Hartmann, and A. O.
  Mooers. 2012. The global diversity of birds in space and time. *Nature*,
 *491*(7424), 444–448.
 [https://doi.org/10.1038/nature](https://doi.org/10.1038/nature) 11631
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 use by army-ant–following birds. *The Auk* 134:247-255. * Chaves-Campos,
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 (2005). Bare-necked umbrellabird (*Cephalopterus glabricollis*) foraging
 at an unusually large assemblage of army-ant-following birds. *The Wilson
 Bulletin* 117:418–420. * Chesser, R. T. (1995). Comparative diets of
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 27:382-390. * Coates-Estrada, R., and A. Estrada (1989). Avian attendance
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 Tuxtlas, Veracruz, Mexico. *Journal of Tropical Ecology* 5:281-292. * Di
 Giacomo, A. S., and A. G. Di Giacomo (2006). Observations of
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Vocal communication in the pale-winged trumpeter (Psophia leucoptera): repertoire, context and functional reference. *Behaviour*, *139*(10), 1331-1359. * Short, L. L., J. F. M. Horne, and A. Bonan (2020). Spot-crowned Barbet (*Capito maculicoronatus*), version 1.0. In Birds of the World (J. del Hoyo, A. Elliott, J. Sargatal, D. A. Christie, and E. de Juana, Editors). Cornell Lab of Ornithology, Ithaca, NY, USA. [https://doi.org/10.2173/bow.spcbar1.01](https://doi.org/10.2173/bow.spcbar1.01). * Stotz, D. F. (1992). Buff-throated Saltator eats army ants. *The Wilson Bulletin* 104:373-374. * Sutton, G. M. (1951). Birds and an ant army in southern Tamaulipas. *The Condor* 53:16-18. * Swartz, M. B. (2001). Bivouac checking, a novel behavior distinguishing obligate from opportunistic species of army-ant-following birds. *The Condor* 103:629-633. * Tórrez, M. A., W. Arendt, and P. Salmeron (2009). 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Populations and local extinctions of birds on Barro Colorado Island, Panama. *Ecological Monographs* 44:153-169. * Willis, E. O. (1979). Comportamento e ecologia da mãe-de-taóca, *Phlegopsis nigromaculata* (D'Orbigny & Lafresnaye)(Aves, Formicariidae). *Revista Brasileira de Biologia* 39:117-159. * Willis, E. O. (1981). *Momotus momota* and *Baryphthengus ruficapillus* (Momotidae) as army ant followers. *Ciência e Cultura* 33:1636-1640. * Willis, E. O. (1982a). The behavior of Black-banded Woodcreepers (*Dendrocolaptes picumnus*). *The Condor* 84:272-285. * Willis, E. O. (1982b). The behavior of Scale-backed Antbirds. *The Wilson Bulletin* 94:447-462. * Willis, E. O. (1982c). The behavior of Black-headed Antbirds (*Percnostola rufifrons*: Formicariidae). *Revista Brasileira de Biologia* 42:233-247. * Willis, E. O. (1982d). Ground-cuckoos (Aves: Cuculidae) as army ant followers. *Revista Brasileira de Biologia* 42:753-756. * Willis, E. O. (1982e). *Notharchus* puffbirds (Aves, Bucconidae) as army ant followers. *Ciência e Cultura* 34:777-782. * Willis, E. O. (1982f). Amazonian *Bucco* and *Monasa* (Bucconidae) as army ant followers. *Ciência e Cultura* 34:782-785. * Willis, E. O. (1982g*). Malacoptila* puffbirds (Bucconidae) as army ant followers. *Ciência e Cultura* 34:924-928. * Willis, E. O. (1982h). The behavior of Red-billed Woodcreepers (*Hylexetastes perrotti*). *Revista Brasileira de Biologia* 42:655-666. * Willis, E. O., and Y. Oniki (1982). Behavior of Fringe-backed Fireeyes (*Pyriglena atra*, Formicariidae): a test case for taxonomy versus conservation. *Revista Brasileira de Biologia* 42:213-223. * Willis, E. O., D. Wechsler, and S. Kistler (1982). *Galbula albirostris* (Aves, Galbulidae), *Trogon rufus* (Trogonidae), and *Electron platyrinchum* (Momotidae) as army ant followers. *Revista Brasileira de Biologia* 42:761-766. * Willis, E. O. (1983a). Tinamous, chickens, guans, rails and trumpeters as army ant followers. *Revista Brasileira de Biologia* 43:19-22. * Willis, E. O. (1983b). *Piaya* cuckoos (Aves, Cuculidae) as army ant followers. *Revista Brasileira de Biologia* 43:29-32. * Willis, E. O. (1983c). Anis (Aves, Cuculidae) as army ant followers. *Revista Brasileira de Biologia* 43:33-36. * Willis, E. O. (1983d). Touracos (Musophagidae), woodpeckers (Picidae), and ovenbirds (Furnariidae) as indirect ant followers. *Ciência e Cultura* 35:804-807. * Willis, E. O. (1983e). Three Dendrocincla woodcreepers (Aves, Dendrocolaptidae) as army ant followers. *Ciência e Cultura* 35:201-204. * Willis, E. O. (1983f). Cis-Andean *Xiphorhynchus* and relatives (Aves, Dendrocolaptidae) as army ant followers. *Revista Brasileira de Biologia* 43:133-142. * Willis, E. O. (1983g). Trans-Andean *Xiphorhynchus* (Aves, Dendrocolaptidae) as army ant followers. *Revista Brasileira de Biologia* 43:125-132. * Willis, E. O. (1983h). Toucans (Ramphastidae) and hornbills (Bucerotidae) as ant followers. *Le Gerfaut* 73:239-242. * Willis, E. O. (1983i). Toucans (Ramphastidae) and Hornbills (Bucerotidae) as ant followers. *Le Gerfaut* 73:239-242. * Willis, E. O. (1983j). Flycatchers, Cotingas, and Drongos (Tyrannidae, Muscicapidae, Cotingidae, and Dicruridae) as ant followers. *Le Gerfaut* 73:265-280. * Willis, E. O. (1983k). Wrens, gnatwrens, rockfowl, babblers and shrikes (Troglodytidae, Polioptilidae, Picathartidae, Timaliidae, and Laniidae) as ant followers. *Le Gerfaut* 73:393-404. * Willis, E. O. (1983l). Jays, mimids, icterids, and bulbuls (Corvidae, Mimidae, Icteridae and Pycnonotidae) as ant followers. *Le Gerfaut* 73:379-392. * Willis, E. O., D. Wechsler, and F. G. Stiles (1983). Forest-falcons, hawks, and a pygmy-owl as ant followers. *Revista Brasileira de Biologia* 43:23-28. * Willis, E. O. (1984a). *Hylophylax*, *Hypocnemoides* and *Myrmoderus* (Aves, Formicariidae) as army ant followers. *Revista Brasileira de Zoologia* 2:159-164. * Willis, E. O. (1984b). Manakins (Aves: Pipridae) as army ant followers. *Ciência e Cultura* 36:817-823. * Willis, E. O. (1984c). *Myrmotherula* antwrens (Aves, Formicariidae) as army ant followers. *Revista Brasileira de Zoologia* 2:153-158. * Willis, E. O. (1984d). *Phlegopsis erythroptera* (Gould, 1855) and relatives (Aves, Formicariidae) as army ant followers. *Revista Brasileira de Zoologia* 2:165-170. * Willis, E. O. (1984e). *Dysithamnus* and *Thamnomanes* (Aves, Formicariidae) as army ant followers. *Papéis Avulsos de Zoologia* 35:183-187. * Willis, E. O. (1984f). Antshrikes (Aves, Formicariidae) as army ant followers. *Papéis Avulsos de Zoologia* 35:177-182. * Willis, E. O. (1984g). Neotropical thrushes (Turdidae) as army ant followers. *Ciência e Cultura* 36:1197-1202. * Willis, E. O. 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