Tree-dwelling ant survey (Hymenoptera, Formicidae) in Mitaraka, French Guiana

Directeur De la publication : Bruno David Président du Muséum national d’Histoire naturelle réDactrice en chef / Editor-in-chiEf : Laure Desutter-Grandcolas assistants De réDaction / AssistAnt Editors : Anne Mabille ([email protected]), Emmanuel Côtez Mise en page / PAgE lAyout : Anne Mabille coMité scientifique / sciEntific boArd : James Carpenter (AMNH, New York, États-Unis) Maria Marta Cigliano (Museo de La Plata, La Plata, Argentine) Henrik Enghoff (NHMD, Copenhague, Danemark) Rafael Marquez (CSIC, Madrid, Espagne) Peter Ng (University of Singapore) Norman I. Platnick (AMNH, New York, États-Unis) Jean-Yves Rasplus (INRA, Montferrier-sur-Lez, France) Jean-François Silvain (IRD, Gif-sur-Yvette, France) Wanda M. Weiner (Polish Academy of Sciences, Cracovie, Pologne) John Wenzel (The Ohio State University, Columbus, États-Unis) couverture / covEr : Plateau forest at Mitaraka (French Guiana) (photograph: Maurice Leponce). In medaillon, Gigantiops destructor (Fabricius, 1804) (Formicinae) (photograph: Maurice Leponce). 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Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2019 ISSN (imprimé / print) : 1280-9551/ ISSN (électronique / electronic) : 1638-9387 163 ZOOSYSTEMA • 2019 • 41 (10) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com urn:lsid:zoobank.org:pub:CCF40E80-D727-409B-AF31-80D93F1D2D55 Leponce M., Delabie J. H. C., Orivel J., Jacquemin J., Calvo Martin M. & Dejean A. 2019. — Tree-dwelling ant survey (Hymenoptera, Formicidae) in Mitaraka, French Guiana, in Touroult J. (ed.), “Our Planet Reviewed” 2015 large-scale biotic survey in Mitaraka, French Guiana. Zoosystema 41 (10): 163-179. https://doi.org/10.5252/zoosystema2019v41a10. http://zoosystema.com/41/10 AbstrAct Ants constitute a substantial part of the arthropod biomass in rainforests. Most studies have focused on ground-dwelling ants, which constitute almost half of the diversity of the ant assemblage. We report here the results of the first survey of tree-dwelling ants in French Guiana on a plateau and in a swamp palm forest (Euterpe oleracea Mart.) in the Mitaraka Mountains. We were interested in seeing the effect of topography and geographic distance on species richness and composition and to gather information Maurice LEPONCE Biodiversity Monitoring & Assessment, Royal Belgian Institute of Natural Sciences (RBINS), Rue Vautier 29, B-1000 Brussels (Belgium) and Evolutionary Biology & Ecology, Université Libre de Bruxelles (ULB), Av. F.D. Roosevelt 50, B-1050 Brussels (Belgium) [email protected] Jacques H. C. DELABIE Laboratório de Mirmecologia, Centro de Pesquisa do Cacau/CEPLAC, Cx.P. 7, 45600-970 Itabuna, Bahia (Brazil) and Departamento de Ciências Agrárias e Ambientais, Universidade Estadual Santa Cruz, Ilhéus, Bahia (Brazil) Jérôme ORIVEL CNRS, UMR EcoFoG Écologie des Forêts de Guyane, AgroParisTech, CIRAD, INRA, Université de Guyane, Université des Antilles, Campus Agronomique, BP 316, F-97379 Kourou Cedex (France) Justine JACQUEMIN Biodiversity Monitoring & Assessment, Royal Belgian Institute of Natural Sciences (RBINS), Rue Vautier 29, B-1000 Brussels (Belgium) Mariano CALVO MARTIN Evolutionary Biology & Ecology, Université Libre de Bruxelles (ULB), Av. F.D. Roosevelt 50, B-1050 Brussels (Belgium) Alain DEJEAN CNRS, UMR EcoFoG, AgroParisTech, Cirad, INRA, Université des Antilles, Université de Guyane, 97310 Kourou (France); and Ecolab, Université de Toulouse, CNRS, INPT, UPS, 31062 Toulouse Cedex 9 (France) Submitted on 27 November 2017 | Accepted on 7 August 2018 | Published on 24 April 2019 Tree-dwelling ant survey (Hymenoptera, Formicidae) in Mitaraka, French Guiana 164 ZOOSYSTEMA • 2019 • 41 (10) Leponce M. et al. on the species distribution on tree trunks. The fauna of Mitaraka was compared with one from a site 350km distant (Petit Saut). In total 105 trees were sampled (30, 30, 45 in the plateau and the swamp forests of Mitaraka, and in Petit Saut plateau forest, respectively). Arboreal ants were attracted using tuna and honey baits spread along a rope reaching an upper branch, except for the palm swamp forest where the baits were only placed at 2 m high. A total of 34, 13 and 22 species were observed in these three respective sites. Six of these species are new records for French Guiana. In Mitaraka Camponotus femoratus (Fabricius, 1804) and Crematogaster levior Longino, 2003 co-occurred on trees (parabiotic association) and were among the most common species, along with Crematogaster tenuicula Forel, 1904 which was found on other trees (species exclusion). The Mitaraka Mountains appeared more species rich and had a species composition distinct from Petit Saut. Topography also influenced ant species composition. Almost half of the species collected by the baitline method were exclusively foraging in the canopy. rÉsUMÉ Inventaire des fourmis arboricoles (Hymenoptera, Formicidae) du Mitaraka, Guyane. Les fourmis constituent une part substantielle de la biomasse d’arthropodes dans les forêts tropicales humides. La plupart des études se sont concentrées sur les fourmis terricoles qui constituent presque la moitié de la diversité de la communauté de fourmis. Nous présentons ici les résultats du premier inventaire de fourmis arboricoles dans une forêt de plateau et une pinotière (Euterpe oleracea Mart.) dans le massif du Mitaraka. Nous nous sommes intéressés à l’effet de la topographie et de l’éloignement géographique sur la richesse et la composition des espèces, ainsi qu’à la distribution verticale des espèces le long des troncs. La faune du Mitaraka a été comparée à celle d’un site éloigné de 350km (Petit Saut). Au total 105 arbres ont été échantillonnées (30, 30, 45 dans les forêts du Mitaraka – plateau et pinotière – et de Petit Saut). Les fourmis arboricoles ont été attirées au moyen d’appâts au thon et au miel, répartis le long d’une corde atteignant une branche sommitale, sauf dans la pinotière où les appâts n’ont été installés qu’à 2m du sol. Un total de respectivement 34, 13 et 22 espèces ont été observées dans les trois sites. Six de ces espèces sont de nouvelles signalisations pour la Guyane. Sur le Mitaraka Camponotus femoratus (Fabricius, 1804) et Crematogaster levior Longino, 2003 coexistent sur les arbres (association parabiotique) et sont parmi les espèces les plus communes avec les Crematogaster tenuicula Forel, 1904 trouvées sur d’autres troncs (exclusion spécifique). Le massif du Mitaraka apparaît plus riche et avec une composition spécifique distincte de Petit Saut. La topographie influence également la composition spécifique. Presque la moitié des espèces récoltées fourragent exclusivement dans la canopée. Mots clÉs Biodiversité, forêts tropicales de plaines, Amazonie, méthode des lignes d’appâts arboricoles, La Planète revisitée, signalisations nouvelles. INTrodUcTIoN Ants are arthropods of major ecological importance in tropical and temperate ecosystems. comprising a single family and with around 15 000described species, their diversity is in the same order of magnitude as birds (Barrowclough etal. 2016; Janicki etal. 2016); as social organisms, they are ubiquitous in tropical terrestrial environments and can be sampled with standardized protocols (Agosti& Alonso 2000). These are the main reasons why they are a manageable arthropod group for biomonitoring (Underwood& Fisher 2006). This contrasts with other arthropod groups such as coleoptera which are much richer in tropical forests (Basset etal. 2012). Many more ant species remain to be discovered in unexplored geographical areas or habitats difficult to sample, such as the soil or the canopy (ryder Wilkie etal. 2007, 2010; Ward 2010). The Amazonian Basin remains one of these little explored regions. Most surveys of local richness have focused on grounddwelling ant communities. For example, in five localities across Guiana, Lapolla etal. (2007) found 230species in the leaf-litter with 38-84species per site. At the Nouragues research Station, French Guiana, from the base to the summit of inselbergs, Groc etal. (2009) found 196species in the leaf litter and put in evidence that ant diversity was influenced by habitat type. In the Brazilian state of Acre, Miranda etal. (2012) sampled 222ant species on the ground with pitfall traps and 115ant species by beating the vegetation. Near Manaus, Brazil, Vasconcelos etal. (2003) also demonstrated the influence of topography, observing more species in river valleys than on forested plateaus. In the same area, at a larger scale (along a 2 000-km-long gradient), Vasconcelos etal. (2010) demonstrated the effect of rainfall and flooding regimes on species turnover. In another survey, using baits on the ground and on the vegetation, Vasconcelos& Vilhena (2006) showed a stratification in the ant assemblage. This stratification was even more thoroughly documented in a study by ryder Wilkie etal. (2010) in Amazonian Ecuador. These authors observed distinct ant assemblages in the soil, on the ground surface, and in the understorey and canopy. of the 489species collected, 282were found only on trees. These ants were obtained by fogging sampling by Terry Erwin who also worked in other parts of Amazonia. In Amazonian Brazil, his fogging campaigns revealed that ants were the most abundant arthropods in trees, representing over one third of KEY WorDs Biodiversity, tropical lowland rainforests, Amazonia, arboreal baitline method, our Planet reviewed, new records. 165 ZOOSYSTEMA • 2019 • 41 (10) Tree-dwelling ants of Mitaraka the collected individuals and biomass (Erwin 1983; Adis etal. 1984). Wilson (1987) found in Erwin’s samples in Peru up to 43ant species from 26genera from a single tree, and Tobin (1997) found 85species from 29genera from two trees and 11associated vines for another site in Peru. The aim of the current study was to complement these studies by: 1) surveying tree-dwelling ants in one of the most remote parts of the Amazon, the Mitaraka Mountains in French Guiana; 2) studying the effect of topography on arboreal ant assemblages; 3) studying the species turnover of tree-dwelling ant assemblages separated by large geographical distances; and 4) gathering information on vertical species distribution along trees. MATErIAL ANd METhodS Study SiteS Mitaraka This species inventory is part of the “our Planet reviewed” (“La Planète revisitée”) French Guiana 2014-2015international biotic survey (Pascal etal. 2015; Touroult etal. 2018). It took place in the Mitaraka mountain range bordering Suriname and Brazil. Two contrasting forest sites were sampled: a plateau (“forêt de plateau”); and a swamp forest with with the palm tree Euterpe oleracea Mart. at the bottom of the valley (“pinotière”). Both were centred on a plot from the diadema (“dIssecting Amazonian diversity by Enhancing a Multiple taxa Approach”) project (http://www.labex-ceba.fr/en/6337-2/). Their latitude and longitude were (2.233°N, –54.444°W) and (2.234°N, –54.448°W), respectively. The diadema project also includes ant samplings on the ground and in the understorey, relying on Winkler extraction, pitfall trapping and vegetation beating. These results have been reported elsewhere (Fichaux 2018). In the plateau forest, 30tree canopies inside a circular plot 30m in radius were sampled (area 0.28ha, Fig.1). In the swamp forest, they were collected from 30trees within a 30× 40m plot (0.12ha, Fig.2). Ants were sampled between the 23rd of February and 8th of March 2017. Petit Saut To compare the faunal composition and diversity of Mitaraka, we used the data from an earlier survey in a plateau forest near the Petit Saut dam (Zone de relâcher, 5.068°N, –52.980°W) between the 13th and 25th of october 2010. A plot of 40× 70m (0.28ha) was delineated there and 45canopy trees Fig. 1. — Plateau forest at Mitaraka: A, location of trees (numbers 1-30) sampled within a circular plot of 30 m radius. The plot was centred on a Diadema plot (axes shown with grey lines); B, baitline around a tree with baits spread at 5 m-intervals along tree trunks. Photo: Maurice Leponce. 6 5 4 29 30 20 180° 195° 210° 225° 240° 255° 270° 285° 300° 315° 330° 335° 345° 0° 15° 30° 45° 60° 75° 90° 105° 120° 135° 150° 165° 28 27 26 3 2 30 7 8 9 10 11 1 12 14 15 18 17 16 19 25 21 23 24 22 20 13 55° 10 m N AB 166 ZOOSYSTEMA • 2019 • 41 (10) Leponce M. et al. were sampled. This plot was centred around a large Azteca cf. chartifex colony to map the spatial extension of its territory. This site was c. 350km away from Mitaraka but part of the same Amazonian forest block. Ant SAmpling We used the arboreal baitline protocol to collect tree-dwelling ants (Leponce& dejean 2011). The baitline protocol consists in putting a rope over the uppermost branch in the canopy using a sling shot (Sherrilltree® Big Shot). Baits are spread every 5m along the rope, from 2m above ground up to the uppermost branch, to detect species vertical stratification. Because the rope forms a loop, baits can be easily collected. Baits consist of a mixture of tuna (in vegetableoil) and honey. These baits represent a source of proteins, lipids and carbohydrates. The mixture is wrapped inside a paper towel and tied to the rope (see Fig.3d, F). Ants dig inside the bait and remain on it even when it is brought back down for inspection. Baits were set in the morning and collected approximately fourhours later, in the afternoon. In the event of heavy rain, baits were removed and replaced the next day. In the swamp forest, the Big Shot could not be used (no firm soil to anchor the pole of the sling shot) and baits were only put 2m above the ground. Atotal of 99, 30 and 146baits were set on the plateau, the swamp and Petit saut forests respectively. This method was designed to collect numerically dominant ants but it also collects part of the other species nesting on trees or at ground level. This method also has the advantage of allowing the vertical distribution of ant species along tree trunks to be assessed. Ant identificAtion Ants were identified on the basis of cEPLAc (commisão Executiva do Plano da Lavoura cacaueira, Itabuna, Brasil) and rBINS (royal Belgian Institute of Natural Sciences) (http://cb.naturalsciences.be/ants/collections/French%20 Guiana%20Ants/) reference collections as well as online resources (AntWeb, AntWiki). Their distribution was checked on AntMaps (AntMaps.org). Specimens collected in Mitaraka and Petit Saut were deposited in the MNhN (Muséum national d’histoire naturelle, Paris) and cEPLAc collections, respectively. Fig. 2. — Palm swamp forest: A, area sampled based on a Diadema plot. Numbers 1-20 indicate the sampling points (not trees) for the Diadema Winkler program, baits were placed 2 m above ground on 30 trees within this area; B, general view of the habitat. Photo: Maurice Leponce. A 5 DH L KGC B AEI FJ 10 13 20 191494 3 2712 17 1611 16 81318 40 m 30 m B 167 ZOOSYSTEMA • 2019 • 41 (10) Tree-dwelling ants of Mitaraka StAtiSticAl AnAlySeS Species rarefaction curves were plotted on the species occurrences data matrices using EstimateS 9.1.0 software (colwell 2016) with 100randomizations of the sampling order without replacement. Trees were considered as sampling units. The number of individuals collected on baits was not taken into account. The rationale is that ants are colonial insects and that it is only the number of colonies that is ecologically meaningful (Longino 2000; Leponce etal. 2004). If an ant species was found on a single or on multiple baits on the same tree its occurrence was counted as 1. To standardize the measurements of species richness between sites, an interpolation to a common number of occurrences (40) was carried out as well as the calculation of the chao2 estimator of species richness. The non-overlapping of 95% confidence intervals constructed from unconditional variance estimators was used as a conservative criterion of statistical difference (colwell 2016). Fig. 3. — Representative ants belonging to the six subfamilies found at Mitaraka and Petit Saut: A, Azteca cf. chartifex Forel, 1878 (Dolichoderinae); B, Ectatomma tuberculatum (Olivier, 1792) (Ectatomminae); C, Gigantiops destructor (Fabricius, 1804) (Formicinae); D, Cephalotes atratus (Linnaeus, 1758) (Myrmicinae); E, Neoponera commutata (Roger, 1860) (Ponerinae); F, Pseudomyrmex faber (Smith, F., 1858) (Pseudomyrmecinae). Photos: Maurice Leponce. Average head width: A, 1.0 mm; B, 2.3 mm; C, 2.5 mm; D, 2.1-4.5 mm; E, 3.2 mm; F, 2.0 mm. A C EF D B 168 ZOOSYSTEMA • 2019 • 41 (10) Leponce M. et al. For the analyses of species co-occurrences designed to reveal positive (i.e., parabiotic association) negative (i.e., exclusion) associations between species, the sampling unit considered was each bait. The independence of the presence of two species was measured with Fisher’s exact-tests. These tests were only conducted on the most frequent species (i.e., present each on at least 10% of the baits). The compositional similarity between two sites was measured with the chao-Jaccard index, which is an appropriate measure in the case of incompletely sampled fauna (chao etal. 2005). ReSultS Species richness A total of 40 tree-dwelling species comprising 15 genera were found at Mitaraka (703 specimens collected). Five species belonging to the genus Pheidole Westwood, 1839 were new records for French Guiana (Table1). The plateau forest was richer than the swamp forest: 34 vs 13 observed species (for 74 and 44 species occurrences, respectively). This difference was significant when considering a standardized species richness: 23.2 ± 6.2 vs 12.5 ± 5.5 species (rarefied richness for 40 occurrences ± 95% confidence interval) (Table1; Fig.4). This is valid even when considering only baits collected near the ground at both sites because the accumulation of species in the plateau forest was similar for groundand canopy-dwelling species (Fig.4). The Mitaraka plateau forest was also richer than the distant forest used as comparison (Petit Saut). This might be due in part to the fact that the Petit Saut plot was largely occupied (19/45 trees) by a single colony of a numerically and aggressively dominant ant (Azteca cf. chartifex Forel, 1896). These Azteca Forel, 1878 were present in massive numbers on baits and excluded most other ant species from their territory. The chao2 estimator indicated that we possibly collected only half of the species present at Mitaraka with a total of 76.6 ± 29.2 and 36.7 ± 20.2 species expected in the plateau and swamp forest, respectively. Taxonomic composition Six ant subfamilies were observed at Mitaraka, namely: Myrmicinae Lepeletier de Saint-Fargeau, 1835 (25 species, 3tribes), Formicinae Latreille, 1809 (7species, 3 tribes), Ectatomminae Emery, 1895 (4 species, 1 tribe), Ponerinae Lepeletier de Saint-Fargeau, 1835 (3 species, 1 tribe), and dolichoderinae Forel, 1878 (1species) (Table1). Some representative species are shown in Figure3. Pseudomyrmecinae Smith, F. 1852 were observed foraging on tree trunks but were not caught at baits. The most species rich genera were Pheidole (10 species), Camponotus Mayr, 1861 (5), Crematogaster Lund, 1831 (5) and Solenopsis Westwood, 1840 (4). Five Pheidole were new records for French Guiana (Pheidole flavens roger, 1863; Pheidole kukrana Wilson, 2003; Pheidole obscurior Forel, 1886; Pheidole pepo Wilson, 2003; Pheidole tristops Wilson, 2003)(Table1). Among additional species found in Petit Saut, Camponotus trapezoideusMayr, 1870, was a new record for French Guiana.The only exotic species observed was Monomorium floricola (Jerdon, 1851).The relative proportion of the different subfamilies, in terms of species numbers, was not different between Mitaraka and Petit Saut (chi-square test, P = 0.8). The faunal similarity between the two Mitaraka sites (chaoJaccard: 0.372) was higher than with the Petit Saut site (0.136 or 0.047 for plateau and swamp comparisons, respectively) (Table1). Among the common species from Mitaraka, only Crematogaster tenuicula Forel, 1904 was found in abundance at Petit Saut as well. Species co-occurrences Seven species were found in both habitat types at Mitaraka, namely (by order of decreasing occurrences on trees): Camponotus femoratus (Fabricius, 1804) (21), Crematogaster tenuicula Forel, 1904 (17), Crematogaster levior Longino, 2003 (14), Nylanderia cf. fulva (Mayr, 1862) (5), Pheidole group flavenssp. 02 roger, 1863 (4), Pheidole transversostriata Mayr, 1887 (3), and Pheidole group flavenssp. 01 roger, 1863 (2). Asignificant negative association was found between the first two species (Fisher’s exact-test; P < 0.05) and marginally between the two Crematogaster (P = 0.07). By contrast a very significant positive association was found between Camponotus femoratus and Crematogaster levior. These two species were particularly abundant in the swamp forest. All other species were found in a single habitat and on less than 15% of the trees. Species vertical distribution These data were only available for the plateau forest at Mitaraka. There, 15 out of 34 species were found up to the canopy (height ≥ 12m) and most of them from the ground to the canopy (between 2 and 22m, n = 10) (Table1). The other 19species were found only above ground (between 2 and 7m) but were species for which only one or two occurrences were noted. Species from Pheidole and Solenopsis were Fig. 4 . — Rarefaction curves representing the species accumulation in the Mitaraka plateau forest (taking into account all baits spread along the trunk or only those near the ground), swamp forest and, by comparison, in the distant site, the plateau forest of “Petit Saut” (baits from the ground to the canopy). Petit Saut 50 40 30 20 10 0 020406080 100 Occurences Species Mitaraka plateau forest, all baits Mitaraka plateau forest, baits near ground only Mitaraka swamp forest, baits near ground only 169 ZOOSYSTEMA • 2019 • 41 (10) Tree-dwelling ants of Mitaraka Table 1. — List of species found at Mitaraka, in the plateau and swamp forests, and in the comparison site Petit Saut. Values represent the number of trees on which each species was observed (with the number of specimens examined under brackets). The minimum and maximum heights at which each species was collected along trees are also provided (in meters above ground). All the species are native to French Guiana, except the exotic Monomorium floricola (Jerdon, 1851), signalled with a *. Full details of examined specimens is provided in Appendix 1. Height Subfamily Tribe Species Plateau Swamp Petit Saut Max Min New records Dolichoderinae – Azteca cf. chartifex Forel, 1878 19(53) 27 2 – –Azteca cf. paraensis Forel, 1904 3(11) 17 2 – Ectatomminae Ectatommini Ectatomma edentatum Roger, 1863 1(1) 17 17 – Ectatomma tuberculatum (Olivier, 1792) 14(20) 17 2 – Gnamptogenys pleurodon (Emery, 1896) 4(38) 17 2 – Gnamptogenys porcata (Emery, 1896) 1(1) 2 2 – Gnamptogenys relicta (Mann, 1916) 1(1) 2 2 – Formicinae Camponotini Camponotus (Myrmaphaenus) sp. 01 1(1) 2 2 – Camponotus (Myrmosphincta) sp. 02 2(8) 2 2 – Camponotus femoratus (Fabricius, 1804) 6(11) 15(36) 17 2 – Camponotus novogranadensis Mayr, 1870 1(1) 7 7 × Camponotus punctulatus andigenus Emery, 1903 1(1) 2 2 – Camponotus trapezoideus Mayr, 1870 1(1) 2 2 – Gigantiopini Gigantiops destructor (Fabricius, 1804) 1(1) 2 2 – Lasiini Nylanderia cf. fulva (Mayr, 1862) 1(3) 4(12) 2 2 – Lasiini Nylanderia sp. 02 1(1) 2 2 – Myrmicinae Attini Cephalotes atratus (Linnaeus, 1758) 2(9) 4(5) 22 2 – Cephalotes marginatus (Fabricius, 1804) 1(1) 4(5) 22 2 – Cephalotes minutus (Fabricius, 1804) 2(2) 7 2 – Ochetomyrmex neopolitus Fernández, 2003 1(4) 7 2 – Pheidole fallax Mayr, 1870 1(9) 12 12 – Pheidole flavens Roger, 1863 4(32) 22 2 × Pheidole group fallax sp. 01 1(8) 2 2 – Pheidole group flavens sp. 01 1(4) 1(5) 2 2 – Pheidole group flavens sp. 02 1(3) 3(11) 12 2 – Pheidole kukrana Wilson, 2003 1(5) 2 2 × Pheidole cf. tobini Wilson, 2003 1(1) 2 2 – Pheidole obscurior Forel, 1886 1(3) 2 2 × Pheidole pepo Wilson, 2003 1(1) 2 2 × Pheidole transversostriata Mayr, 1887 2(23) 1(12) 7 2 – Pheidole group tristis sp. 01 1(1) 2 2 – Pheidole group tristis sp. 02 2(2) 2 2 – Pheidole tristops Wilson, 2003 1(3) 2 2 × Strumigenys sp. 01 1(1) 2 2 – Wasmannia auropunctata (Roger, 1863) 1(7) 2 2 – Wasmannia rochai Forel, 1912 1(3) 2(4) 12 2 – Crematogastrini Crematogaster brasiliensis Mayr, 1878 16(22) 22 2 – Crematogaster curvispinosa Mayr, 1862 3(37) 2 2 – Crematogaster erecta Mayr, 1866 8(21) 22 2 – Crematogaster levior Longino, 2003 6(50) 8(27) 17 2 – Crematogaster limata Smith F., 1858 1(4) 5(7) 12 2 – Crematogaster longispina Emery, 1890 1(10) 2 2 – Crematogaster tenuicula Forel, 1904 13(141) 4(22) 9(10) 22 2 – Solenopsidini Megalomyrmex leoninus Forel, 1885 1(6) 2 2 – Monomorium floricola (Jerdon, 1851)* 1(1) 7 7 – Rogeria subarmata (Kempf, 1961) 1(3) 2 2 – Solenopsis sp. 01 4(77) 17 2 – Solenopsis sp. 02 1(6) 2 2 – Solenopsis sp. 03 2(7) 17 17 – Solenopsis sp. 04 3(41) 17 2 – Solenopsis sp. 05 3(3) 7 2 – Solenopsis sp. 06 4(5) 2 2 – Ponerinae Ponerini Neoponera apicalis (Latreille, 1802) 1(1) 2 2 – Neoponera commutata (Roger, 1860) 2(4) 2 2 – Neoponera villosa (Fabricius, 1804) 1(1) 2 2 – Odontomachus haematodus (Linnaeus, 1758) 1(2) 2 2 – Pseudomyrmecinae Pseudomyrmex faber (Smith F., 1858) 2(2) 2 2 – Species 34 13 22 Occurrences (abundance) on trees 74 (523) 44 (180) 102 (172) Trees sampled 30 30 45 Rarefied richness (40 occurrences) (± 95% CI) 23.2 ± 6.2 12.5 ± 5.5 15.3 ± 4.1 Chao2 (± 95%CI) 76.6 ± 29.2 36.7 ± 20.2 29.8 ± 6.2 176 ZOOSYSTEMA • 2019 • 41 (10) Leponce M. et al. Pheidole group flavenssp. 01 mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, swamp forest, 2.234°N, –54.448°W, 5-7.III.2015. 53194-d26-5/3/2015 (5); 53214-c22-7/3/2015 (4). diStRiBution.— French Guiana. Pheidole group flavenssp.02 mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, swamp forest, 2.234°N, –54.448°W, 2-5.III.2015. 50943-c30-2/3/2015 (3); 53151-d14-5/3/2015 (2); 53165-d305/3/2015 (3); 53165-d40-5/3/2015 (6). diStRiBution.— French Guiana. Pheidole kukrana Wilson, 2003 Pheidole kukrana Wilson, 2003: 310. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 2.III.2015. 50942-c30-2/3/2015 (5). diStRiBution.— Neotropical region: costa rica, French Guiana (new record), Guadeloupe, Guatemala, Nicaragua (type locality), Venezuela. Pheidole cf. tobini Wilson, 2003 Pheidole tobini Wilson, 2003: 356. mAteRiAl exAmined.— French Guiana. Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 13.X.2010. 42894-PS21-13/10/2010 (1). diStRiBution.— Native in French Guiana. Pheidole group tristis (Smith, F. 1858) Pheidole group tristis sp.01 mAteRiAl exAmined.— French Guiana. Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 20.X.2010. 43082-PS34-20/10/2010 (1). diStRiBution.— French Guiana. Pheidole group tristis sp. 02 mAteRiAl exAmined.— French Guiana. Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 14-16.X.2010. 42914-PS13-14/10/2010 (1); 42993-PS38-16/10/2010 (1). diStRiBution.— Native in French Guiana. Pheidole obscurior Forel, 1886 Pheidole obscurior Forel, 1886: 7. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 7.III.2015. 53212-c11-7/3/2015 (3). diStRiBution.— Neotropical region: Argentina, Barbados, Brazil, colombia, costa rica, dominican republic, El Salvador, French Guiana (new record), Guatemala (type locality), honduras, Mexico, Nicaragua, Paraguay, Puerto rico, Trinidad and Tobago, Venezuela. Pheidole pepo Wilson, 2003 Pheidole pepo Wilson, 2003: 730. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 2.III.2015. 53168-c26-2/3/2015 (1). diStRiBution.— Neotropical region: colombia (type locality), French Guiana (new record). Pheidole transversostriata Mayr, 1887 Pheidole transversostriata Mayr, 1887: 584. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, swamp forest, 2.234°N, –54.448°W, 27.II5.III.2015. 50952-c12-27/2/2015 (20); 50953-c12-27/2/2015 (2); 50970c09-3/3/2015 (1); 53152-d15-5/3/2015 (12). d iStRiBution .— Neotropical region: Barbados, Brazil, colombia, French Guiana, Guyana (type locality), Suriname, Trinidad and Tobago, Venezuela. Pheidole tristops Wilson, 2003 Pheidole tristops Wilson, 2003: 770. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 3.III.2015. 53135-c04-3/3/2015 (3). diStRiBution.— Neotropical region: colombia (type locality), French Guiana (new record). Genus Strumigenys Smith F., 1860 Strumigenyssp. 01 mAteRiAl exAmined.— French Guiana. Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 13.X.2010. 42877-PS04-13/10/2010 (1). diStRiBution.— French Guiana. 177 ZOOSYSTEMA • 2019 • 41 (10) Tree-dwelling ants of Mitaraka Wasmannia auropunctata (roger, 1863) Tetramorium auropunctatumroger, 1863: 182. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 3.III.2015. 53131-c21-3/3/2015 (7). diStRiBution.— Afrotropical region: cameroun, Gabon, Sierra Leone. Australasian region: Australia, New caledonia. IndoAustralian region: hawaii, Solomon Islands, Vanuatu. Nearctic region: canada, United States. Neotropical region: Antigua and Barbuda, Argentina, Aruba, Barbados, Belize, Bermuda, Bolivia, Brazil, colombia, costa rica, cuba (type locality), dominican republic, Ecuador, French Guiana, Galapagos Islands, Grenada, Guadeloupe, Guatemala, haiti, honduras, Lesser Antilles, Mexico, Netherlands Antilles, Panama, Paraguay, Peru, Puerto rico, Saint Lucia, Uruguay, Venezuela. Palaearctic region: Israel, Spain. Wasmannia rochai Forel, 1912 Wasmannia rochai Forel, 1912: 1. 50958-c27-2/3/2015 (3) mAteRiAl exAmined.— French Guiana. Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 13-19.X.2010. 42897-PS23-13/10/2010 (1); 42925-PS24-14/10/2010 (1); 43020PS24-19/10/2010 (1); 43021-PS24-19/10/2010 (1). diStRiBution.— Neotropical region: Argentina, Brazil (type locality), costa rica, Ecuador, French Guiana, Guatemala, Guyana, Mexico, Panama, Suriname, Trinidad and Tobago, Venezuela. Tribe cRemAtogAStRini Lund, 1831 Genus Crematogaster Lund, 1831 Crematogaster brasiliensis Mayr, 1878 Crematogaster brasiliensis Mayr, 1878: 875. mAteRiAl exAmined.— French Guiana. Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 13-20.X.2010. 42875-PS02-13/10/2010 (1); 42876-PS03-13/10/2010 (1); 42879-PS06-13/10/2010 (1); 42884-PS11-13/10/2010 (1); 42886-PS13-13/10/2010 (1); 42899-PS26-13/10/2010 (1); 42904-PS03-14/10/2010 (1); 42905-PS04-14/10/2010 (1); 42907-PS06-14/10/2010 (1); 42944-PS06-15/10/2010 (1); 42951-PS09-15/10/2010 (1); 42988-PS33-16/10/2010 (1); 42990-PS35-16/10/2010 (1); 42992-PS37-16/10/2010 (1); 42995-PS40-16/10/2010 (1); 43024-PS26-19/10/2010 (1); 43032-PS43-20/10/2010 (1); 43033-PS42-20/10/2010 (1); 43075PS38-20/10/2010 (1); 43076-PS38-20/10/2010 (1); 43082-PS3420/10/2010 (1); 43093-PS43-20/10/2010 (1). diStRiBution.— Neotropical region: Bolivia, Brazil (type locality), colombia, costa rica, Ecuador, Frecnh Guiana, Guatemala, honduras, Mexico, Nicaragua, Panama, Peru, Trinidad and Tobago. Crematogaster curvispinosa Mayr, 1862 Crematogaster curvispinosa Mayr, 1862: 768. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, swamp forest, 2.234°N, –54.448°W, 5.III.2015. 53146-d08-5/3/2015 (2); 53155-d18-5/3/2015 (28); 53156d19-5/3/2015 (7) diStRiBution.— Neotropical region: Argentina, Barbados, Bolivia, Brazil (type locality), colombia, costa rica, Ecuador, French Guiana, Galapagos Islands, Grenada, Guadeloupe, Guatemala, Guyana, honduras, Lesser Antilles, Martinique, Mexico, Nicaragua, Panama, Paraguay, Peru, Suriname, Uruguay, Venezuela. Crematogaster erecta Mayr, 1866 Crematogaster erecta Mayr, 1866: 902. mAteRiAl exAmined.— French Guiana. Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 13-21.X.2010. 42877-PS04-13/10/2010 (1); 42903-PS02-14/10/2010 (1); 42905-PS04-14/10/2010 (1); 42935-PS02-15/10/2010 (1); 42936-PS02-15/10/2010 (1); 42937-PS02-15/10/2010 (1); 42939-PS04-15/10/2010 (1); 42940-PS04-15/10/2010 (1); 42947-PS07-15/10/2010 (1); 42948-PS09-15/10/2010 (1); 42949-PS09-15/10/2010 (1); 42950-PS09-15/10/2010 (1); 43016-PS21-19/10/2010 (1); 43051-PS21-20/10/2010 (1); 43052-PS21-21/10/2010 (1); 43072-PS40-20/10/2010 (1); 43074-PS39-20/10/2010 (1); 43093-PS43-20/10/2010 (1); 43099PS40-20/10/2010 (1); 43100-PS40-20/10/2010 (1); 93101-PS3920/10/2010 (1). diStRiBution.— Neotropical region: Brazil, colombia, costa rica (type locality), Ecuador, French Guiana, Guyana, Mexico, Panama, Peru, Suriname. Crematogaster levior Longino, 2003 Crematogaster levior Longino, 2003: 132. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, swamp forest, 2.234°N, –54.448°W, 27.II-7.III.2015. 50950-c14-27/2/2015 (13); 50957-c08-2/3/2015 (3); 53129-c193/3/2015 (10); 53130-c22-7/3/2015 (7); 53134-c25-7/3/2015 (16); 53141-d01-5/3/2015 (6); 53143-d04-5/3/2015 (4); 53147d09-5/3/2015 (2); 53154-d17-5/3/2015 (5); 53161-d23-5/3/2015 (2); 53162-d22-5/3/2015 (4); 53171-c28-2/3/2015 (1); 53191d27-5/3/2015 (3); 53192-d25-5/3/2015 (1). diStRiBution.— Neotropical region: Bolivia, Brazil (type locality), colombia, Ecuador, French Guiana, Guyana, Peru, Suriname, Venezuela. Crematogaster limata Smith F., 1858 Crematogaster limatusSmith F., 1858: 139. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 2.III.2015, Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 16-20.X.2010. 53137-c05-2/3/2015 (4). 42987-PS32-16/10/2010 (1); 42999-PS44-16/10/2010 (1); 43000-PS45-16/10/2010 (1); 43002-PS47-16/10/2010 (1); 43023-PS27-19/10/2010 (1); 43031-PS44-20/10/2010 (1); 43063PS27-19/10/2010 (1). diStRiBution.— Neotropical region: Bolivia, Brazil (type locality), colombia, costa rica, Ecuador, Guatemala, Guyana, honduras, Mexico, Nicaragua, Panama, Peru, Trinidad and Tobago, Venezuela. 178 ZOOSYSTEMA • 2019 • 41 (10) Leponce M. et al. Crematogaster longispina Emery, 1890 Crematogaster longispina Emery, 1890: 53. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 7.III.2015. 53139-c09-7/3/2015 (10). d iStRiBution .— Neotropical region: colombia, costa rica (type locality), Ecuador, French Guiana, Guyana, Nicaragua, Peru, Suriname. Crematogaster tenuicula Forel, 1904 Crematogaster tenuicula Forel, 1904: 36. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, swamp forest, 2.234°N, –54.448°W, 27.II-7.III.2015, Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 16-20.X.2010. 50944-c01-27/2/2015 (28); 50963-c01-1/3/2015 (7); 50964-c021/3/2015 (5); 50965-c02-1/3/2015 (28); 50969-c08-2/3/2015 (4); 50973-c03-1/3/2015 (1); 53128-c18-3/3/2015 (1); 53136-c042/3/2015 (1); 53138-c06-2/3/2015 (1); 53142-d02-5/3/2015 (4); 53145-d07-5/3/2015 (3); 53153-d16-5/3/2015 (10); 53159-d295/3/2015 (5); 53169-c26-2/3/2015 (8); 53170-c27-2/3/2015 (8); 53172-c28-2/3/2015 (2); 53175-c29-2/3/2015 (1); 53208-c247/3/2015 (32); 53211-c25-7/3/2015 (14). 42881-PS08-13/10/2010 (1); 42898-PS24-13/10/2010 (1); 42901-PS28-13/10/2010 (1); 42927-PS26-14/10/2010 (1); 42985-PS30-16/10/2010 (1); 42989-PS34-16/10/2010 (1); 42993PS38-16/10/2010 (1); 42997-PS42-16/10/2010 (1); 43078-PS3520/10/2010 (1); 43091-PS30-20/10/2010 (1). diStRiBution.— Neotropical region: costa rica to Amazonian Brazil (including French Guiana), Bolivia. Tribe SolenopSidini Forel, 1893 Genus Megalomyrmex Forel, 1885 Megalomyrmex leoninus Forel, 1885 Megalomyrmex leoninus Forel, 1885: 372. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 27.II.2015. 50949-c15-27/2/2015 (6). diStRiBution.— Neotropical region: Brazil, colombia (type locality), Ecuador, French Guiana, Guyana, Suriname, Trinidad and Tobago, Venezuela. Genus Monomorium Mayr, 1855 Monomorium floricola (Jerdon, 1851) Atta floricolaJerdon, 1851: 107. mAteRiAl exAmined.— French Guiana. Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 15.X.2010. 42931-PS01-15/10/2010 (1). diStRiBution.— Afrotropical region: cameroun, comoros, Ghana, Nigeria, Togo, United republic of Tanzania. Australasian region: Australia, New caledonia, Norfolk Island. Indo-Australian region: Borneo, cook Islands, Fiji, French Polynesia, Guam, hawaii, Indonesia, Kiribati, Krakatau Islands, Malaysia, Marshall Islands, Micronesia (Federated States of), New Guinea, Niue, Northern Mariana Islands, Palau, Philippines, Samoa, Solomon Islands, Tokelau, Tonga, Vanuatu, Wallis and Futuna Islands. Malagasy region: Mauritius, Mayotte, réunion, Seychelles. Nearctic region: United States. Neotropical region: Anguilla, Bahamas, Barbados, Brazil, cayman Islands, chile, colombia, costa rica, cuba, dominican republic, Ecuador, El Salvador, French Guiana, Galapagos Islands, Greater Antilles, Grenada, Guadeloupe, Guatemala, Guyana, haiti, Lesser Antilles, Mexico, Paraguay, Puerto rico, Suriname, Trinidad and Tobago. oriental region: India (type locality), Laos, Nicobar Island, SriLanka, Thailand, Vietnam. Palaearctic region: china, Japan, republic of Korea, United Kingdom of Great Britain and Northern Ireland. Exotic to French Guiana. Genus Rogeria Emery, 1894 Rogeria subarmata (Kempf, 1961) Irogera subarmataKempf, 1961: 438. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 2.III.2015. 53171-c28-2/3/2015 (3). diStRiBution.— Neotropical region: Brazil (type locality), Ecuador, French Guiana, Venezuela. Genus Solenopsis Westwood, 1840 Solenopsissp. 01 mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 27.II2.III.2015. 53171-c28-2/3/2015 (3); 50951-c14-27/2/2015 (15); 50954-c1227/2/2015 (46); 50955-c12-27/2/2015 (10); 50962-c08-2/3/2015 (1); 50968-c06-2/3/2015 (5). diStRiBution.— French Guiana. Solenopsissp. 02 mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, swamp forest, 2.234°N, –54.448°W, 5.III.2015. 53151-d14-5/3/2015 (6). diStRiBution.— French Guiana. Solenopsissp. 03 mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 3.III.2015. 50980-c16-3/3/2015 (1); 53132-c21-3/3/2015 (6). diStRiBution.— French Guiana. Solenopsissp. 04 mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 2-7.III.2015. 179 ZOOSYSTEMA • 2019 • 41 (10) Tree-dwelling ants of Mitaraka 50960-c21-3/3/2015 (8); 53175-c29-2/3/2015 (30); 53206-c11-7/3/2015 (2); 53210-c21-3/3/2015 (1). diStRiBution.— French Guiana. Solenopsissp. 05 mAteRiAl exAmined.— French Guiana. Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 16-20.X.2010. 42986-PS31-16/10/2010 (1); 43028-PS45-19/10/2010 (1); 43081-PS29-20/10/2010 (1). diStRiBution.— French Guiana. Solenopsissp. 06 mAteRiAl exAmined.— French Guiana. Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 13-20.X.2010. 42898-PS25-13/10/2010 (1); 42912-PS11-14/10/2010 (1); 42926PS25-14/10/2010 (1); 42984-PS29-16/10/2010 (1); 43087-PS3120/10/2010 (1). diStRiBution.— French Guiana. Subfamily poneRinAe, Lepeletier de Saint-Fargeau, 1835 Tribe poneRini, Lepeletier de Saint-Fargeau, 1835 Genus Neoponera Emery, 1901 Neoponera apicalis (Latreille, 1802) Formica apicalisLatreille, 1802: 204. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, swamp forest, 2.234°N, –54.448°W, 5.III.2015. 53160-d24-5/3/2015 (1). diStRiBution.— Neotropical region: Belize, Bolivia, Brazil, colombia, costa rica, Ecuador, French Guiana, Guatemala, Guyana, honduras, Mexico, Nicaragua, Panama, Peru, Suriname, Trinidad and Tobago, Venezuela. Neoponera villosa (Fabricius, 1804) Formica villosaFabricius, 1804: 409. mAteRiAl exAmined.— French Guiana. Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 13.X.2010. 42897-PS23-13/10/2010 (1). diStRiBution.— Nearctic region: United States. Neotropical region: Argentina, Belize, Bolivia, Brazil, colombia, costa rica, Ecuador, El Salvador, French Guiana, Guatemala, Guyana, honduras, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto rico, Suriname, Trinidad and Tobago, Venezuela. Neoponera commutata (roger, 1860) Ponera commutataroger, 1860: 311. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, plateau forest, 2.233°N, –54.444°W, 2.III.2015. 53167-c01-2/3/2015 (2); 53167-c26-2/3/2015 (2). diStRiBution.— Neotropical region: Brazil, colombia, Ecuador, French Guiana, Guyana, Paraguay, Peru, Suriname. Genus Odontomachus Latreille, 1804 Odontomachus haematodus (Linnaeus, 1758) Formica haematodaLinnaeus, 1758: 582. mAteRiAl exAmined.— French Guiana. Mitaraka, “our Planet reviewed”, swamp forest, 2.234°N, –54.448°W, 5.III.2015. 53149-d11-5/3/2015 (2). d iStRiBution .— Nearctic region: United States. Neotropical region: Argentina, Bahamas, Bolivia, Brazil, colombia, costa rica, Ecuador, French Guiana, Greater Antilles, Grenada, Guadeloupe, Guatemala, Guyana, honduras, Lesser Antilles, Mexico, Nicaragua, Panama, Paraguay, Peru, Suriname, Trinidad and Tobago, Venezuela. Palaearctic region: china. Subfamily pSeudomyRmecinAe Smith F., 1852 Genus Pseudomyrmex Lund, 1831 Pseudomyrmex faber (Smith F., 1858) Pseudomyrma faberSmith F., 1858: 157. mAteRiAl exAmined.— French Guiana. Petit Saut dam, Zone de relâcher, 5.068°N, –52.980°W, 13-15.X.2010. 42875-PS02-13/10/2010 (1); 42930-PS01-15/10/2010 (1). diStRiBution.— Neotropical region: Brazil (type locality), colombia, costa rica, Ecuador, French Guiana, Guyana, Panama, Suriname.