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 ('averagemass') is the average mass of the species in grams. The eight column ('Beak.LengthCulmen') 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 11631

Ant-following Level

• Batcheller, H. J. (2017). Interspecific information use by army-ant–following birds. The Auk 134:247-255.
• Chaves-Campos, J. (2003). Localization of army-ant swarms by ant-following birds on the Caribbean slope of Costa Rica: following the vocalization of antbirds to find the swarms. Ornitología Neotropical 14:289-294.
• Chaves-Campos, J. (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 obligate ant-following birds at a site in northern Bolivia. Biotropica 27:382-390.
• Coates-Estrada, R., and A. Estrada (1989). Avian attendance and foraging at army-ant swarms in the tropical rain forest of Los Tuxtlas, Veracruz, Mexico. Journal of Tropical Ecology 5:281-292.
• Di Giacomo, A. S., and A. G. Di Giacomo (2006). Observations of Strange‐tailed Tyrants (Alectrurus risora) and other grassland birds following army-ants and armadillos. Journal of Field Ornithology 77:266-268.
• Dobbs, R. C., and P. R. Martin (1998). Migrant bird participation at an army-ant swarm in montane Jalisco, Mexico. Wilson Bulletin 110:293-295.
• Driver, R. J., S. DeLeon, and S. O'Donnell (2018). Novel observation of a raptor, Collared Forest-falcon (Micrastur semitorquatus), depredating a fleeing snake at an army ant (Eciton burchellii parvispinum) raid front. The Wilson Journal of Ornithology 130:792-796.
• Faria, C., and M. Rodrigues (2009). Birds and army-ants in a fragment of the Atlantic Forest of Brazil. Journal of Field Ornithology 80:328-335.
• Gochfeld, M., and G. Tudor (1978). Ant-following birds in South American subtropical forests. Wilson Bulletin 90:139-141.
• Greene, E., D. Wilcove, and M. McFarland (1984). Observations of birds at an army-ant swarm in Guerrero, Mexico. The Condor 86:92-93.
• Greeney, H.F. 2012. Antpittas and worm-feeders: a match made by evolution? Evidence for a possible commensal foraging relationship between antpittas (Grallariidae) and mammals. Neotropical Biology and Conservation 7: 140-143.
• Greeney, H. F. (2020). Amazonian Trogon (Trogon ramonianus), version 1.0. In Birds of the World (S. M. Billerman, B. K. Keeney, P. G. Rodewald, and T. S. Schulenberg, Editors). Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bow.viotro3.01.
• Haemig, P. D. (1989). Brown Jays as army ant followers. The Condor 91:1008-1009.
• Hardy, J. W. (1974). Jays as army ant followers. The Condor 76:102-103.
• Hilty, S. L. (1974). Notes on birds at swarms of army-ants in the highlands of Colombia. The Wilson Bulletin 86:479-481.
• Johnson, R. A. (1954). The behavior of birds attending army-ant raids on Barro Colorado Island, Panama Canal Zone. Proceedings of the Linnaean Society of New York 65:41-70.
• Karr, J. R. (1971). Ecological, behavioral, and distributional notes on some central Panama birds. The Condor 73:107-111.
• Karubian, J., and L. Carrasco (2008). Home range and habitat preferences of the Banded Ground-cuckoo (Neomorphus radiolosus). The Wilson Journal of Ornithology 120:205-209.
• Klein, B. C., and R. O. Bierregaard (1988). Movement and calling behavior of the Lined Forest-Falcon (Micrastur gilvicollis) in the Brazilian Amazon. The Condor 90:497-499.
• Kumar, A., and S. O'Donnell (2007). Fragmentation and elevation effects on bird–army-ant interactions in neotropical montane forest of Costa Rica. Journal of Tropical Ecology 23:581-590.
• Lane, D. F., T. Valqui, J. Alvarez, J. Armenta, and K. Eckhardt (2006). The rediscovery and natural history of the White‐masked Antbird (Pithys castaneus). The Wilson Journal of Ornithology 118:13-22.
• Maldonado-Coelho, M., and R. Durães (2003). The black-goggled tanager (Trichothraupis melanops): an occasional kleptoparasite in mixed-species bird flocks and ant swarms of Southeastern Brazil. Ornitología Neotropical 14:397-403.
• Mays, N. M. (1985). Ants and foraging behavior of the Collared Forest-Falcon. The Wilson Bulletin 97:231-232.
• Nieto-R., M., and J.D. Ramírez. 2006. Notas sobre aves de tierras altas que siguen marchas de hormigas arrieras para su alimentación, en la Reserva Natural Río Blanco, Manizales, Caldas. Boletín de la Sociedad Antioqueña de Ornitología 16: 59-66.
• O'Donnell, S., A. Kumar, and C. Logan (2010). Army-ant raid attendance and bivouac-checking behavior by Neotropical montane forest birds. The Wilson Journal of Ornithology 122:503-512.
• O'Donnell, S., A. Kumar, and C. J. Logan (2014). Do Nearctic migrant birds compete with residents at army-ant raids? A geographic and seasonal analysis. The Wilson Journal of Ornithology 126:474-487.
• O'Donnell, S. (2017). Evidence for facilitation among avian army‐ant attendants: specialization and species associations across elevations. Biotropica 49:665-674.
• Oniki, Y. (1971). Wandering interspecific flocks in relation to ant-following birds at Belem, Brazil. The Condor 73:372-374.
• Oniki, Y. (1972). Studies of the guild of ant-following birds at Belem, Brazil. Acta Amazonica 2:59-79.
• Oniki, Y., and E. O. Willis (1972). Studies of ant-following birds north of the eastern Amazon. Acta Amazonica 2:127-151.
• Pizo, M. A., and A. S. Melo (2010). Attendance and co-occurrence of birds following army-ants in the Atlantic rain forest. The Condor 112:571-578.
• Rasmussen, P. C., and N. Collar (2020). Black-streaked Puffbird (Malacoptila fulvogularis), 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.blspuf1.01.
• Rasmussen, P. C., N. Collar, and G. M. Kirwan (2020a). Guianan Puffbird (Notharchus macrorhynchos), 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.guipuf1.01.
• Rasmussen, P. C., N. Collar, and G. M. Kirwan (2020b). Buff-bellied Puffbird (Notharchus swainsoni), 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.bubpuf1.01.
• Rios, M., G. Londoño, and L. Biancucci (2008). Notes on birds that follow army ants in the northern Andes. Ornitología Neotropical 19:137-142.
• Rivera, J. H. V., D. Ayala, and C. A. Haas (2003). Home-range size, habitat use, and reproduction of the Ivory-billed Woodcreeper (Xiphorhynchus flavigaster) in dry forest of western Mexico. Journal of Field Ornithology 74:141-151.
• Roberts, D. L., R. J. Cooper, and L. J. Petit (2000). Flock characteristics of ant‐following birds in premontane moist forest and coffee agroecosystems. Ecological Applications 10:1414-1425.
• Roberts, D. L. (2016). Two species of tanagers (Passeriformes: Thraupidae) forage on army ant workers (Eciton burchellii) carrying immature paper wasps. The Wilson Journal of Ornithology 128:653-656.
• Rodrigues, M., C. G. Machado, and M. Galetti (1994). Association of the Black-goggled Tanager (Trichothraupis melanops) with flushers. Biotropica 26:472-475.
• Sainz-Borgo, C. (2015). Consumo de obreras Eciton burchellii (Hymenoptera: Formicidae) por varias especies de aves en condiciones urbanas. Ecotrópicos 28:38-42.
• Sazima, I., and G. B. D'Angelo (2015). Association of birds with social insects: a summary in Southeastern Brazil. Iheringia: Série Zoologia 105:333-338.
• Seddon, N., Alvarez, A., & Tobias, J. (2002). 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.
• 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). Aves hormigueras en bosque seco del Pacífico de Nicaragua: uso de hábitat y comportamiento parasítico. Zeledonia 13:1-9.
• Vallely, A. C. (2001). Foraging at army ant swarms by fifty bird species in the highlands of Costa Rica. Ornitología Neotropical 12:271-275.
• Willis, E. O. (1960). A study of the foraging behavior of two species of ant tanagers. The Auk 77:150-170.
• Willis, E. O. (1966a). Interspecific competition and the foraging behavior of plain‐brown woodcreepers. Ecology 47:667-672.
• Willis, E. O. (1966b). The role of migrant birds at swarms of army ants. Living Bird 5:187-231.
• Willis, E. O. (1966c). Ecology and behavior of the Crested Ant-Tanager. The Condor 68:56-71.
• Willis, E. O. (1967). The behavior of Bicolored Antbirds. University of California Publications in Zoology 79:1-132.
• Willis, E. O. (1968a). Taxonomy and behavior of Pale-faced Antbirds. The Auk 85:253-264.
• Willis, E. O. (1968b). Studies of the behavior of Lunulated and Salvin's Antbirds. The Condor 70:128-148.
• Willis, E. O. (1969). On the behavior of five species of Rhegmatorhina, ant-following antbirds of the Amazon basin. The Wilson Bulletin 81:363-395.
• Willis, E. O. (1972a). The behavior of Plain-brown Woodcreepers, Dendrocincla fuliginosa. The Wilson Bulletin 84:377-420.
• Willis, E. O. (1972b). The behavior of Spotted Antbirds. American Ornithologists Union Monographs 10:1-162.
• Willis, E. O. (1972c). Taxonomy, ecology and behavior of the Sooty Ant-Tanager and other ant-tanagers. American Museum Novitates 2480:1-38.
• Willis, E. O., and Y. Oniki (1972). Ecology and nesting behavior of the Chestnut-backed Antbird (Myrmeciza exsul). The Condor 74:87-98.
• Willis, E. O. (1973). The behavior of Ocellated Antbirds. Smithsonian Contributions to Zoology 144:1-57.
• Willis, E. O. (1974). 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. (1985a). Cercomacra and related antbirds (Aves, Formicariidae) as army ant followers. Revista Brasileira de Zoologia 2:427-432.
• Willis, E. O. (1985b). Antthrushes, antpittas, and gnateaters (Aves, Formicariidae) as army ant followers. Revista Brasileira de Biologia 2:443-448.
• Willis, E. O. (1985c). Myrmeciza and related antbirds (Aves, Formicariidae) as army ant followers. Revista Brasileira de Zoologia 2:433-442.
• Willis, E. O. (1986a). Vireos, wood warblers and warblers as ant followers. Le Gerfaut 76:177-186.
• Willis, E. O. (1986b). Tanagers, finches and weavers as ant followers. Le Gerfaut 76:307-316.
• Willis, E.O. (1988). "Behavioral notes, breeding records, and range extensions for Colombian birds". Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales. 16 (63): 137–150.
• Willis, E. O., and Y. Oniki (1992). As aves e as formigas de correição. Boletim do Museu
  Paraense Emílio Goeldi, Zoologia 8:123-150.
• Willis, E. O., and Y. Oniki (1993). On Dendrocincla tyrannina: morphology, behavior and conservation of a shy lek-type insectivore. Caldasia 18:131-140.
• Willson, S. K. (2004). Obligate army-ant-following birds: a study of ecology, spatial movement patterns, and behavior in Amazonian Peru. Ornithological Monographs 55:1-67.
• Zimmer, K. J., and S. L. Hilty (1997). Avifauna of a locality in the upper Orinoco drainage of Amazonas, Venezuela. Ornithological Monographs 48:865-885.

Morphological Data

• Tobias, J. A., Sheard, C., Pigot, A. L., Devenish, A. J., Yang, J., Sayol, F., ... & Schleuning, M. (2022). AVONET: morphological, ecological and geographical data for all birds. Ecology Letters, 25(3), 581-597.

Elevation Minimum and Maximum Data

• BirdLife International. 2023. IUCN Red List for birds. Downloaded from http://www.birdlife.org.
• Stotz, D. F., T. Parker III, and D. K. Moskovits. 1996. Neotropical birds: ecology and conservation. University of Chicago Press.
• The Cornell Lab of Ornithology. n.d.. Birds of the world. https://birdsoftheworld.org/bow/home

Vertical Foraging Strata

• Wilman, H., J. Belmaker, J. Simpson, C. de la Rosa, M. M. Rivadeneira, and W. Jetz. 2014. EltonTraits 1.0: Species-level foraging attributes of the world’s birds and mammals. Ecology, 95(7), 2027–2027. https://doi.org/10.1890/13-1917.1
• Wilman, H., J. Belmaker, J. Simpson, C. de la Rosa, M. M. Rivadeneira, and W. Jetz. 2016. EltonTraits 1.0: Species-level foraging attributes of the world's birds and mammals. Wiley. Collection. https://doi.org/10.6084/m9.figshare.c.3306933.v1