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Social Structure of Gnamptogenys Bisulca (Formicidae: Ectatomminae) in Tropical Forests

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Social Structure of Gnamptogenys Bisulca (Formicidae: Ectatomminae) in Tropical Forests Insectes Sociaux https://doi.org/10.1007/s00040-019-00716-w Insectes Sociaux RESEARCH ARTICLE Social structure of Gnamptogenys bisulca (Formicidae: Ectatomminae) in tropical forests D. M. Urcuqui1 · J. Herrera‑Rangel1 · C. Poteaux2 · I. Armbrecht1 Received: 9 February 2019 / Revised: 27 May 2019 / Accepted: 29 June 2019 © International Union for the Study of Social Insects (IUSSI) 2019 Abstract The study of the social organization of ants may help to understand why some species are able to persist in natural forests after fragmentation and anthropogenic disturbances. The Neotropical ant Gnamptogenys bisulca has shown to be a biologi- cal indicator of habitat quality in Andean montane forests and we proposed to explain this characteristic by investigating its social structure and the spatial distribution of colonies at fne scale. In eight 100 m2 plots located in four forests in the western Colombian Andes, the position of G. bisulca colonies was recorded for spatial distribution and their social composi- tion described: ergatoid females were found in most of the nests, isolated or together with normal queens. As ergatoids and queens were dissected to examine their reproductive status, it appeared that apparent polygyny represented, in fact, efective monogyny, with other potential reproductive females being unmated or inhibited. In the few cases of mixed colonies, a queen was always at the head of the colony. Local spatial distribution was random and did not ft the hypothesis of nesting by fs- sion, as generally observed in the case of the presence of ergatoids. However, the social structure was signifcantly diferent according to the site, the most preserved forest showing no or rare ergatoids (but queens) compared to the other sites. Our results indicated that the presence of ergatoids in G. bisulca may help colonies to adapt to slightly disturbed habitat contexts. Keywords Ergatoids · Monogyny · Polygyny · Tropical montane forest Introduction absence of wings in queens because this variation of the queen phenotype is generally linked to their dispersal tactics While the division of labor is a key factor in social hymenop- (Molet et al. 2009): mainly large winged queens are devoted terans explaining their ecological success, ants in particular, to far-away dispersion (through independent colony found- present huge social plasticity, with diferences not only in ing, ICF) while wingless or reduced-size queens disperse the number of queens per colony (polygyny), but also their with nestmate workers to found a new colony by budding size and morphology may vary among species, populations or fssion (dependent colony founding, DCF). Many recent and occasionally even among colonies within a population reviews have pointed out that wing dimorphism in the queen (Bourke and Franks 1995; Crozier and Pamilo 1996; Rosset is relatively common across all lineages of ants (Molet et al. and Chapuisat 2007; Heinze 2008; Lenoir et al. 2011). Many 2012; Peeters 2012; Peeters and Adams 2016; Johnson and authors have focused on the polymorphism of the presence/ Overson 2018). However, the morphology of reproductive females is also associated with the presence of several typi- Electronic supplementary material The online version of this cal queen traits linked to their fecundity (e.g., presence of article (https ://doi.org/10.1007/s0004 0-019-00716 -w) contains a spermatheca, high number of ocelli, complex structured supplementary material, which is available to authorized users. thorax, large gaster and developed ovaries) that are reduced or absent in non-reproductive castes. * I. Armbrecht [email protected] According to the novel model of developmental evolution in ants proposed by Molet et al. (2012), rare intermediate 1 Department of Biology, Universidad del Valle, Calle 13 morphotypes, such as intercaste individuals, have acciden- 100-00. Ed. E20, Of. 3025, Cali, Colombia tally deviated from their normal developmental pathways 2 Laboratoire d’Ethologie Expérimentale et Comparée due to environmental or genetic perturbations. Thus, they EA 4443, Université Paris 13, Sorbonne Paris Cité, present mosaics of winged queens and worker characters. 93430 Villetaneuse, France Vol.:(0123456789)1 3 D. M. Urcuqui et al. The morphology of intercastes is reported to be highly varia- most abundant species in the Winkler extraction of leaf lit- ble in several species (e.g., Temnothorax nylanderi, Plateaux ter. Inside both riparian and secondary forests, the species 1970; Molet et al. 2012; Johnson and Overson 2018) while is among the most abundant, accounting for approximately the morphology of ergatoid (worker-like apterous queens, 40% of captures (Jiménez-Carmona et al. 2015). As expected Amor et al. 2011) can vary into distinct categories or not, for being a deep-forest species, it is almost absent in severely according to the species considered (review in Peeters 2012). transformed habitats, such as crops or pasturelands, even The study of ergatoid queens is particularly interesting to in ecologically restored ones (Donoso and Ramón 2009; understand the evolutionary signifcance of these alternative Abadía et al. 2010; Zabala et al. 2013; Chacón de Ulloa and phenotypes, since they present a simplifed thorax similar to Abadía 2014; Herrera-Rangel et al. 2015). that of workers—as a consequence of their absence of wing Despite its ecological importance as a forest bioindicator muscles—but are fully reproductive females (Peeters and (Herrera-Rangel et al. 2015), the natural history and social Adams 2016). They can occur together with alate queens structure of G. bisulca is poorly known. Therefore, this in the same colony, or replace them, but rarely occur with study sought to explore the social composition (number of brachypterous queens, other short-winged morphotypes that individuals, castes, morphology of reproductive females) in cannot fy (Johnson and Overson 2018). colonies of G. bisulca to understand the role of ergatoids, Among the ants, the Ectatomminae (Hymenoptera: For- both in their contribution to reproduction and to spatial micidae) are relatively specialized hunter ants, whose natu- structuration of the population (DCF, polydomy). To test ral history and ecology has been largely unknown (Lattke for ecological infuences on the social organization in this 1995). However, several studies have pointed out the vari- species, we investigated the colony structure and distribution ability in the social organization of these ants: microgynes of G. bisulca in four diferent populations from natural mon- in Ectatomma ruidum (Lachaud et al. 1999; Lenoir et al. tane forests in Colombia difering in the degree of habitat 2011), gamergates due to the presence of a spermatheca disturbance: one site is located in a protected natural park in workers of Gnamptogenys menadensis and G. striatula while the rest of them are immersed in a matrix of unshaded (Gobin et al. 1998, 2001; Blatrix and Jaisson 2000; Lom- cofee crops. melen et al. 2006) but also the presence of both ergatoids and gamergates in some populations of G. striatula (Blatrix and Jaisson 2000; Giraud et al. 2001; Kaptein et al. 2003). Gnamptogenys bisulca Kempf and Brown is a species Materials and methods known from Mexico, Costa Rica, Ecuador and Colombia (García-Martínez et al. 2015). In this last country, it has been Study area and sampling for colony distribution recorded in most of the western states, at altitudes between 1000 and 2000 m.a.s.l., and from humid to dry tropical for- Sampling was carried out in several villages of La Celia ests (Lattke et al. 2008). The species, which was described Municipality, Risaralda Department, west-center in Colom- from leaf litter samples in Colombia (Kempf and Brown bia (Fig. 1; Online Resource 1). The studied landscape pre- 1968) nests in rotten wood, mainly in small twigs on the for- sented a broken relief with rugged slopes, and heterogeneous est foor (Jiménez-Carmona 2015). Their colonies are small, sight, containing towns, forest relicts (~ 24% overall), open of approximately 30 individuals, and the female reproduc- sun cofee (~ 65%), but also open cattle pastures, plantain, tive caste is composed of queens and sometimes ergatoids. maize and bean plantations among other crops (Zabala et al. While colonies frequently, but not always, present ergatoids 2013). The study was carried out inside forests located in (approximately 12% of nests, Jiménez-Carmona et al. 2015), four independent villages with a minimum of 4 km separa- their role has never been investigated. tion distance: La Secreta, El Brillante, El Tambo and La This variability in social organization may help to under- Polonia (Fig. 1; Online Resource 1). La Secreta forest, is stand why some species are able to persist in natural habi- located in Verdum Regional Natural Park, and is considered tats (usually forests) after severe fragmentation and anthro- the most conserved forest. The remaining forests were sur- pogenic disturbances. In this way, G. bisulca Kempf and rounded by sun cofee plantations in fragmented landscapes. Brown is an appropriate model because it has been described The areas of all forests studied for spatial distribution were as a biological indicator of forested habitat (Abadía et al. calculated using ArcGIS software (Version 10.3). All of 2010; Herrera-Rangel et al. 2015). Most information about them shared similar climatic conditions, within the natural G. bisulca has been gathered from sampling studies deal- zone of the Very Humid Premontane Forest, with
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