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The Arboreal Ants of a Neotropical Rain Forest Show High Species Density and Comprise One Third of the Ant Fauna

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Received: 16 April 2019 | Revised: 7 October 2019 | Accepted: 7 January 2020 DOI: 10.1111/btp.12772 ORIGINAL ARTICLE The arboreal ants of a Neotropical rain forest show high species density and comprise one third of the ant fauna John T. Longino1 | Robert K. Colwell2,3 1Department of Biology, The University of Utah, Salt Lake City, UT, USA Abstract 2Department of Ecology and Evolutionary In tropical rain forests, the ant community can be divided into ground and arboreal Biology, University of Connecticut, Storrs, faunas. Here, we report a thorough sampling of the arboreal ant fauna of La Selva CT, USA 3University of Colorado Museum of Natural Biological Station, a Neotropical rain forest site. Forty-five canopy fogging samples History, Boulder, CO, USA were centered around large trees. Individual samples harbored an average of 35 ant Correspondence species, with up to 55 species in a single sample. The fogging samples yielded 163 John T. Longino, Department of Biology, observed species total, out of a statistically estimated 199 species. We found no rela- The University of Utah, Salt Lake City, UT 84112, USA. tionship between within-sample ant richness and focal tree species, nor were the ant Email: [email protected] faunas of nearby trees more similar to each other than the faunas of widely spaced Funding information trees. Species density was high, and beta diversity was low: A single column of veg- National Science Foundation, Grant/Award etation typically harbors at least a fifth of the entire arboreal ant fauna. Considering Number: DEB-1354739 the entire fauna, based on 23,326 species occurrence records using a wide variety Associate Editor: Emilio Bruna of collecting methods, 182 of 539 observed species (196 of 605, estimated statisti- Handling Editor: Emilio Bruna cally) were entirely arboreal. The arboreal ant fauna is thus about a third of the total La Selva ant fauna, a robust result because inventory completeness was similar for ground and arboreal ants. The taxonomic history of discovery of the species that make up the La Selva fauna reveals no disproportionately large pool of undiscovered ant species in the canopy. The "last biotic frontier" for tropical ants has been the rot- ten wood, leaf litter, and soil of the forest floor. Abstract in Spanish is available with online material. KEYWORDS arthropods, canopy, Costa Rica, fogging, formicidae 1 | INTRODUCTION the ground and canopy communities overlap very little (Longino & Colwell, 1997; Ryder Wilkie, Mertl, & Traniello, 2010; Yanoviak & TROPICAL RAIN FORESTS ARE RENOWNED FOR THEIR COMPLEX STRUCTURE AND Kaspari, 2000). One set is entirely ground-based, nesting in the soil, HIGH DIVERSITY. Ants are a dominant biotic element in rain forests leaf litter, or rotten wood of the shaded forest floor and foraging (Davidson, Cook, Snelling, & Chua, 2003; Fittkau & Klinge, 1973; there or within a few meters of the ground. Another set is arboreal, Floren, Biun, & Linsenmair, 2002; Stork, 1988), and a growing body nesting and foraging mostly or entirely in the vegetation. Only a few of research documents the abundance, diversity, behavior, and hab- species routinely occur in both habitats. itat specificity of rain forest ants. Ant faunas are vertically strati- Arboreality is a derived trait in ants that has evolved repeatedly fied with respect to their nesting and foraging preferences, and in many ant lineages (Dejean, Corbara, Orivel, & Leponce, 2007; © 2020 The Association for Tropical Biology and Conservation Biotropica. 2020;00:1–11. wileyonlinelibrary.com/journal/btp | 1 2 | LONGINO AND COLWELL Nelson, Ree, & Moreau, 2018). A few of the arboreal lineages assessment of the arboreal fauna based on all collecting methods. have radiated spectacularly, forming the bulk of the abundance We use these results to estimate the total richness of arboreal ants and diversity of arboreal ants. The development of arboreal ant and to compare the richness of arboreal ants and ground ants. We communities is primarily a tropical phenomenon; the abundance of test for an effect of focal tree species on sample richness and species arboreal ants is orders of magnitude greater in the tropics than in composition, and we test for an effect of spatial proximity on species the temperate zone (Floren, Wetzel, & Staab, 2014). Tropical biolo- composition. The taxonomy of the arboreal ant species is carefully gists have focused their research on assessing (a) the total richness considered, and species are linked to specimens with unique spec- of arboreal ants (LePonce et al., 2019; Majer, 1990; Wilson, 1987; imen identifiers in museum collections, insuring that the study can Yanoviak, Fisher, & Alonso, 2007); (b) how arboreal ant richness serve as a reference benchmark for future work. compares to richness of ground ants (Floren et al., 2014; Ryder Wilkie et al., 2010; Yanoviak & Kaspari, 2000); (c) whether arbo- real ants form "mosaics" of dominant and subdominant species 2 | METHODS (Dejean et al., 2016, 2019, 2007, 2018, 2015; Floren & Linsenmair, 2000; Jackson, 1984; LePonce et al., 2019; Leston, 1978; Majer & 2.1 | Study site and fogging methods Camer-Pesci, 1991; Room, 1971); and (d) whether ant communities are affected by habitat, tree species, tree size, or connectivity via The study was carried out at La Selva Biological Station, Heredia lianas (Adams, Schnitzer, & Yanoviak, 2017; Floren & Linsenmair, Province, Costa Rica (McDade, Bawa, Hespenheide, & Hartshorn, 1997, 2005; Klimes, Fibich, Idigel, & Rimandai, 2015; Klimes et al., 1993), from 1993 to 2000. Mean annual rainfall is approximately 2012; Osorio-Pérez, Barberena-Arias, & Aide, 2007; Philip, Fayle, 4 m. The habitat is a mosaic of mature lowland rain forest, second & Yusah, 2018; Ribas, Schoereder, Pic, & Soares, 2003; Schonberg, growth forest of various ages, and abandoned pastures. The study Longino, Nadkarni, Yanoviak, & Gering, 2004; Schulz & Wagner, area comprised approximately 1,500 ha, ranging from 50 m to 150 m 2002). in elevation. La Selva Biological Station in Costa Rica, one of the best-stud- Canopy fogging was carried out using the general procedures ied tropical rain forests, has been particularly well sampled for discussed by Erwin (1983), Adis, Lubin, and Montgomery (1984), and ants (Longino & Colwell, 1997; Longino, Colwell, & Coddington, Stork (1988). A total of 45 fogging samples were acquired (Figure 1, 2002). Ant sampling at La Selva was carried out over a period of Table 1). For 11 samples, the focal tree was Pentaclethra macroloba 13 years, with a combination of canopy fogging, Malaise traps, light (Willd.) O. Ktze. (Fabaceae), the most common tree species at La traps, Berlese samples, Winkler samples of sifted litter, baiting, and Selva. For 12 samples, the focal tree was Virola koschnyi Warb. hand-collecting. Some of the data have been previously published (Myristicaceae), a species of intermediate abundance. The remaining in the context of inventory efficiency (Longino & Colwell, 1997) and 22 samples targeted focal trees from a wide range of species and richness estimation of the entire ant fauna (Longino et al., 2002). The plant families. A stratified sampling design was used for the first 18 previous publications used only a portion of the fogging data. Here, fogging samples, with trees fogged in six sets of three closely spaced we present an analysis of the entire fogging dataset, along with an individuals (Figure 1). In each group of three samples, one tree was FIGURE 1 Spatial distribution of fogging samples at La Selva Biological Station, Costa Rica. The first 18 samples are numbered, showing the triplet structure of the sampling. Latitudes and longitudes of each sample are in Table 1. The grid cells are 1 km2 LONGINO AND COLWELL TABLE 1 Fogging samples. See text for explanation of processing methods Funnels Processing Sample code Date Latitude Longitude Species processed method Sobs S40 iChao2 FVK/01 14-Jan−93 10.43278 −84.00894 Virola koschnyi (Myristicaceae) 40 1 28 28 44.31 FOT/02 5-Mar−93 10.43486 −84.0127 Carapa guianensis (Meliaceae) 20 1 50 57.88 65.04 FPM/03 6-Mar−93 10.43454 −84.01195 Pentaclethra macroloba (Fabaceae) 17 1 33 43.50 50.66 FOT/04 5-May−93 10.41274 −84.00395 Conceveiba pleiostemona (Euphorbiaceae) 10 1 30 37.97 38.10 FPM/05 6-May−93 10.41317 −84.00379 Pentaclethra macroloba (Fabaceae) 10 1 32 45.71 50.05 FVK/06 7-May−93 10.41406 −84.00432 Virola koschnyi (Myristicaceae) 10 1 28 43.56 46.09 FOT/07 3-Jul−93 10.42411 −84.00792 Goethalsia meiantha (Tiliaceae) 10 1 20 34.30 39.31 FPM/08 4-Jul−93 10.42461 −84.00746 Pentaclethra macroloba (Fabaceae) 11 1 44 52.72 54.58 FVK/09 5-Jul−93 10.42414 −84.0073 Virola koschnyi (Myristicaceae) 10 1 48 63.44 68.42 FOT/10 3-Sep−93 10.42582 −84.0133 Tapirira guianensis (Anacardiaceae) 10 1 27 42.86 46.75 FVK/11 4-Sep−93 10.42663 −84.01268 Virola koschnyi (Myristicaceae) 10 1 49 54.39 54.55 FPM/12 5-Sep−93 10.42685 −84.01129 Pentaclethra macroloba (Fabaceae) 11 1 8 14.88 18.00 FPM/13 6-Nov−93 10.42543 −84.00488 Pentaclethra macroloba (Fabaceae) 10 1 51 64.60 67.38 FOT/14 8-Nov−93 10.42519 −84.00512 Sacoglottis trichogyna (Humiriaceae) 10 1 21 28.07 28.99 FVK/15 9-Nov−93 10.42487 −84.0048 Virola koschnyi (Myristicaceae) 10 1 15 25.73 28.52 FOT/16 5-Jan−94 10.43012 −84.01293 Vitex cooperi (Verbenaceae) 11 1 33 46.51 48.14 FVK/17 7-Jan−94 10.43076 −84.01245 Virola koschnyi (Myristicaceae) 10 1 21 24.66 24.68 FPM/18 8-Jan−94 10.4315 −84.01283 Pentaclethra macroloba (Fabaceae) 10 1 26 34.78 36.24 FVK/19 8-Oct−94 10.4203 −84.01137 Virola koschnyi
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  • Hymenoptera: Formicidae) Community Over 25 Years

    Hymenoptera: Formicidae) Community Over 25 Years

    Changes in an invaded Florida ant (Hymenoptera: Formicidae) community over 25 years Leo Ohyama1,*, Joshua R. King1, and Benjamin M. Gochnour2 Abstract Exotic invasive species are one of the most widespread and common agents of change in ecosystems worldwide. Here, we are focused on community- level changes associated with the appearance and persistence of exotic species in an ecosystem over more than 2 decades. We combined datasets of Florida’s ant (Hymenoptera: Formicidae) community from Wekiwa Springs State Park, Florida, USA, spanning 25 yr, which included 3 sampling events in several distinct upland ecosystems. Species accumulation curves, non-parametric species estimators, community similarity indices, and ratios of exotic to native ants were used to assess sampling effort, including patterns of diversity, and changes in community composition. Our data showed that the ant community of Wekiwa Springs has at least 4 exotic species present in all of the ecosystems sampled within the park. These upland eco- systems are accumulating exotics slowly and, with one exception, there is no clear signal that the exotic species are displacing native species. The likely exception is the invasive fire ant,Solenopsis invicta Buren (Hymenoptera: Formicidae), causing local extinction of the native fire Solenopsisant geminata (Fabricius) (Hymenoptera: Formicidae). Continued long-term monitoring efforts are necessary to understand how these exotics may impact native communities in the future. Key Words: displacement; fire; fire ants; ecosystem management; invasion; persistence Resumen Las especies exóticas invasoras son uno de los agentes comunes más extendidos de cambio en los ecosistemas de todo el mundo. Aquí, nos enfocamos sobre los cambios a nivel comunitario asociados con la aparición y persistencia de especies exóticas en un ecosistema durante más de 2 décadas.
  • Tropical Ant Communities Are in Long-Term Equilibrium

    Tropical Ant Communities Are in Long-Term Equilibrium

    Ecological Indicators 83 (2017) 515–523 Contents lists available at ScienceDirect Ecological Indicators j ournal homepage: www.elsevier.com/locate/ecolind Original Articles Tropical ant communities are in long-term equilibrium David A. Donoso Instituto de Ciencias Biológicas, Escuela Politécnica Nacional, Av. Ladrón de Guevara E11-253, Quito, Ecuador a r t i c l e i n f o a b s t r a c t Article history: Communities change with time. Studying long-term change in community structure permits deeper Received 13 June 2016 understanding of community dynamics, and allows us to forecast community responses to perturbations Received in revised form 10 March 2017 at local (e.g. fire, secondary succession) and global (e.g. desertification, global warming) spatial scales. Accepted 13 March 2017 Monitoring efforts exploring the temporal dynamics of indicator taxa are therefore a critical part of Available online 23 March 2017 conservation agendas. Here, the temporal dynamics of the Otongachi leaf litter ant community, occurring in a cloud forest in coastal Ecuador, were explored. By sampling this community six times over eleven Keywords: years, I assessed how the ant fauna caught by Winkler traps (more diverse and cryptic fauna) and caught Andes cloud forests Monitoring by pitfall traps (larger, more mobile fauna) changed over time. The Otongachi leaf litter ant community Otongachi was dynamic. Although species richness in the community remained constant, temporal turnover of Rank abundance species was high: on average, 51% of the ant species in Winkler traps, and 56% of those in pitfall traps, Temporal turnover were replaced with other ant species from one year to the other.