DOI: 10.13102/sociobiology.v67i4.5558Sociobiology 67(4): 492-500 (December, 2020) Open access journal: http://periodicos.uefs.br/ojs/index.php/sociobiology ISSN: 0361-6525 Introduction Tropical forest canopies are known to support a high animal diversity. Particularly arthropods can be very diverse and abundant (Klimes et al., 2012). Ants account for 20 to 60 % of total arthropod biomass of tropical forest canopy invertebrates (Floren et al., 2014), and therefore, represent one of the most abundant and ecologically important animal groups in tropical regions (Hölldobler & Wilson, 1990; Lach et al., 2010). Ants are also widely considered as keystone species due to the important ecological role they play in many ecosystems (Del Toro et al., 2012). In tropical forest canopy, ants often exercise a high predation pressure and thus significantly influence structure and dynamics of arboreal arthropod communities (Philpott & Armbrecht, 2006). Ant are also Abstract Ants constitute an important part of arboreal arthropod biomass in rainforests. Nevertheless, there are only a few methods which permit a rapid assessment of these insects in the canopy layer. This study aims at evaluating the efficiency of a new variant type of pitfall trap i.e. “the funnel trap”, to sample arboreal ants in a secondary and gallery forest in Lamto reserve (Côte d’Ivoire). This method was compared to standard arboreal pitfall trap and beating. In total, the 3 methods yielded 7072 ant workers belonging to 43 species, 14 genera and 5 subfamilies. Tree beating recorded the highest ant’s numerical abundance (3670 workers), with 27 species, 12 genera and 3 subfamilies followed by the “funnel trap” that yielded 2800 ant workers, with 23 species belonging to 12 genera and 5 subfamilies. Finally, arboreal pitfall traps caught the lowest individual with 602 ant workers from 20 species belonging to 9 genera and 3 subfamilies. The composition of species which are caught by arboreal pitfall trap and “funnel trap” was similar at 53 percent. Tree beating showed a distinct species composition compared to arboreal pitfall trap and “funnel trap”. The “funnel trap” could be a fast and efficient way to quickly assess ant-biodiversity in forest canopies and agroecosystems as it looks like a non- destructive sampling method. Sociobiology An international journal on social insects CD Yodé1,2, K Dosso2, LMM Kouakou2 , K Yeo2, W Dekoninck3, S Konate2, PK Kouassi1 Article History Edited by Gilberto M. M. Santos, UEFS, Brazil Received 06 July 2020 Initial acceptance 07 September 2020 Final acceptance 16 November 2020 Publication date 28 December 2020 Keywords Pitfall trap, “funnel” trap, Lamto reserve, biodiversity, habitat structure, tree canopy. Corresponding author Christine Dakélé YODÉ Felix Houphouët Boigny University Research Station in Ecology of Lamto Scientific Reserve BP 28 N’Douci, Côte d’Ivoire. E-Mail: christineyode@gmail.com used as indicator taxa for ecological surveys because they are relatively easy to sample in soil litter (Alonso & Agosti, 2000). Despite this importance, the arboreal ant community remains poorly known for some region in tropics, mainly because the difficulty for scientists to find appropriate sampling method to sample either at three meters above the ground. For example, methods to sample arboreal ants such as the spikes- and-belt method, single-rope technique, baited pitfall traps and so forth, are methods for which one needs an appropriate logistic to access to the canopy and for the installation of all sampling equipments (Basset et al., 1997). Nevertheless, few techniques such as the canopy fogging and beating do not require a lot of equipment in the canopy (Castaño-Meneses, 2014). Currently, pitfall traps (including its variants) are often used for arboreal ant sampling (Powell et al., 2011; Chapin & Smith, 2019). 1 - Felix Houphouët Boigny University, (UFR Biosciences), Zoology-Biology of Animal Laboratory, Abidjan, Côte d’Ivoire 2 - Nangui Abrogoua University, UFR des Sciences de la Nature (UFR SN), Research Station in Ecology of Lamto, Abidjan, Côte d’Ivoire 3 - Royal Belgian Institute of Natural Sciences, OD Taxonomy and Phylogeny, Brussels, Belgium RESEARCH ARTICLE - ANTS Evaluating Efficiency of Different Sampling Methods for Arboreal Ants (Hymenoptera: Formicidae) in A West African Forest-Savanna Mosaic Sociobiology 67(4): 492-500 (December, 2020) 493 Although, pitfall trapping is a well-recognized ant sampling technique, an important difficulty remains the installation of these traps above three meters high in the canopy. Usually this installation requires climbing which is difficult and risky. Hence, to sample canopy biota rigorous inventory techniques need to be developed. These techniques should be simple, fast and allow researchers to optimize their sampling in the canopy (Yusah et al., 2012; Yusah et al., 2018; Leponce et al., 2019). Here, this study reports the results of arboreal ant diversity surveys using different sampling methods in a tropical forest-savanna mosaic habitat at Lamto scientific reserve (Côte d’Ivoire). Overall, it aims at establishing a database on arboreal ant species richness in Côte d’Ivoire. Indeed, the arboreal ant community remains poorly known and the only existing studies go back to the seventies (Delage-Darchen, 1970, 1971, 1972, 1973, 1974; Levieux, 1976). This study aims at testing the efficiency of a new variant of pitfall trap, the “funnel trap” to collect arboreal ants in secondary and gallery forest of Lamto scientific reserve. This trap was compared with arboreal baited pitfall trap and beating of tree leaves, by analysing the differences in numbers of ant workers, species richness and species composition. Methods Study site The study was conducted in Lamto Scientific Reserve, located in central Côte d’Ivoire at 6°13’/6°25’ N and 4°97’/5°01’ W. The annual precipitation range is 1000–1500 mm/year while the mean monthly temperature is about 27°C. Lamto Scientific reserve contains a great heterogenous vegetation (Abbadie et al., 2006) characterized by a forest- savanna mosaic habitat. The study was carried out in the gallery forest located at the border of Bandama river, and the secondary forest that resulted from an experimental bush fire exclusion since 1962 (Abbadie et al., 2006; Gnaoré et al., 2018). Sampling design and identification Ants were sampled in three 100 × 50 m plots in gallery forests and secondary forest. On each plot, 20 trees with a circumference at breast height ≥ 32 cm were examined. Overall samplings were carried out on 120 trees spread over 6 plots in all. On each plot, three sampling methods were used to collect arboreal ants: canopy beating, arboreal pitfall trap (Agosti & Alonso, 2000; Underwood & Fisher 2006; Yeo et al ., 2013) and the “funnel trap”, a modified variant of the arboreal pitfall trap (Fig 1). Fig 1. “Funnel trap” model. Arboreal pitfall trap: These traps consisted of plastic cups (7.5 cm in diameter, 10.5 cm deep) with water, detergent and baited with tuna (Ribas et al., 2003). They were placed on the central axis of the trees at least at 5 m above the ground thanks to a point. They were installed using a ladder. Pitfall traps remained in service on the trees during 48 h. The “funnel trap”: The latter was made from empty bottles of 33 cl of mineral water or soda, rope and a bamboo shaft. Bottles were cut in half at 8 cm from the top edge. Next, the upper part of the bottle was placed upside down in the second part in order to have a funnel. These two parts were attached to their joints with a transparent adhesive tape. Finally, two small holes were made at 1cm of the top edge of the bottle to insert a fine nylon thread whose length corresponds to the height at which the trap can be attached to the tree. To prevent the thread from mixing, it was previously wind up on small pieces of wood 7 cm. Finally, tuna bait was placed inside the bottle with a little water and was hoisted at 5 meters into the trees using a bamboo of at least 5m height (Fig 2). The traps remained in service on during 48 hours. The sampling was carried out on 120 trees. CD Yodé, K Dosso, LMM Kouakou, K Yeo, W Dekoninck, S Konate, PK Kouassi – The “funnel trap”: a new sampling method for arboreal ants494 Canopy beating: This method consisted in collecting the ants by beating the foliage of the lowest branches of the trees (between 2m and 3m high). The foliage of the trees was beaten in two sequences of 5 beats. Each sequence was followed by the gathering of the ants that fell on the canvas. In total the foliage was beaten ten times at the same point. Also, it had 20 collection points on a plot, and therefore 120 collection points in all. Fig 2. Installation of “funnel trap” in tree canopy. 10 0 m 50 m Fig 3. Scheme of the sampling plot in the Lamto Scientific Reserve, Côte d’Ivoire. Arboreal pitfall traps and funnel traps were installed on the same trees. As for the beating, they were generally carried out on the same trees or in thickets directly connected to the trees in which the traps were installed (Fig 3). Ants were identified to genus level using the guide of the genera to Fisher and Bolton (2016) with a Leica MZ6 microscope. At species level, the keys of Bolton (1980, 1982 and 1987); Rigato, 2016), and reliable digital keys (antweb.org) were used. When species-level identification was impossible, distinct specimens were sorted according to morphospecies. Morphospecies were numbered according to the ant reference collection for Côte d’Ivoire at Lamto ecological research station. The specimens were added to Lamto ecological research station collection. Data analysis Samples of each plot were pooled to obtain a total of 120 samples for each sampling method. Taxonomic structure (subfamily), species richness, abundance, and species composition were compared for each sampling method. The Chao 2 index species richness estimator was calculated to extrapolate the species richness from our data. Sample coverages were determined to estimate sampling efficiency based on different sampling method using EstimateS v.9.1.0. Comparative analysis was carried out on the species richness and numerical abundance of the ants collected by the different trap types. Kruskall-Wallis and Man-Whitney pairwise comparison tests were used to test the differences across the trap types using species richness and numerical abundance. The comparison of species composition was conducted calculating Jaccard similarity index. Results General results Overall, the three combined methods yielded 7072 ant workers belonging to 43 species, 14 genera and 5 subfamilies. The subfamilies were Dolichoderinae, Myrmicinae, Formicinae, Ponerinea and Pseudomyrmicinae. All five subfamilies were collected with the funnel trap, whereas only three out of five subfamilies were collected with arboreal pitfall traps and beating, respectively (Fig 4). Sampling efficiency Table 1 shows that the sample coverage varied between 71 and 93 %, illustrating that all three methods were suitable to investigate arboreal ant community. Observed species Observed species Estimated species (Chao 2) Sampling coverage Unique species Doubletons Arboreal pitfall 20 21,49 93% 4 3 Funnel trap 23 32,26 71% 8 2 Beating 27 31,17 86% 7 4 Table 1. Numbers of species caught by the trapping methods during study. Sociobiology 67(4): 492-500 (December, 2020) 495 accumulation curves evolved towards asymptotic lines for all sampling methods. However, the Chao 2 estimated species accumulation curves for “funnel trap” and beating increased steadily with sampling size (Fig 5). In addition, the tree beating method collected the highest number of species (27 species), followed by funnel trap (23 species) and arboreal pitfall trap (20 species). Funnel traps caught more unique species (8 species) followed by beating (7 species) and arboreal pitfall (4 species) (Table 1). Species richness and diversity Tree beating recorded the highest number of ant species (27 species) with 3670 workers, followed by the “funnel trap” (23 species) for 2800 workers. Arboreal pitfall traps caught the lowest number of ant species (20 species) and workers (602). The mean ant species number found in traps differed significantly between the 3 sampling methods (Kruskall- Fig 4. Ant species distribution within the different subfamilies encountered for each sampling method. Fig 5. Species accumulation curves of the three sampling methods. Wallis: X 2 = 89.66; df = 2; p = 0.0001). Tree beating method recorded the highest mean number of ant species (2.59 species/trap) followed by “funnel trap” method (1.63 species/ trap) and arboreal pitfall trap method (0.98 species/trap). The Man-Whitney pairwise comparison test showed that “funnel trap” caught more ant species than arboreal pitfall trap (U = 4670; p < 0.001). On the other hand, tree beating method caught more ant species than “funnel trap” (U = 4216; p < 0.0001) and arboreal pitfall trap (U = 2335; p < 0.001) (Such as Fig 6). The three sampling methods yielded a high diversity, but the value of the Simpson diversity index was higher for the beating method (0.89), followed by that of the funnel trap (0.86) and finally the arboreal pitfall trap (0.84). On the other hand, evenness values were low, with E=0.34 for the beating and E=0.36 for both funnel trap and arboreal pitfall trap, respectively (Table 2). CD Yodé, K Dosso, LMM Kouakou, K Yeo, W Dekoninck, S Konate, PK Kouassi – The “funnel trap”: a new sampling method for arboreal ants496 Ant abundance and species composition Overall, ant abundance (individuals) varied significantly among each sampling method (ANOVA of Kruskal-Wallis test: X 2 = 85.23; df = 2; p = 0.0001; Such as Fig 7). On average, the tree beating method (29.93 individuals/trap) and “funnel trap” (23.33 individuals/trap) caught the highest numbers of ant workers, whilst arboreal pitfall trap caught 5.02 individuals/trap. Mann-Whitney pairwise comparison test indicated that the funnel trap caught more ant workers than the arboreal pitfall trap (U = 3931; p = 0.001). However, the tree beating method caught more ant workers (individuals) than the funnel trap (U = 5686; p = 0.004), and arboreal pitfall trap (U = 2307; p = 0.001). Globally, beating yielded a different species composition that funnel trap and arboreal pitfall trap. Otherwise, Jaccard similarity index showed that the arboreal pitfall trap and funnel trap had approximately a similar species composition with a similarity percentage at 53% of shared species. Tree beating shared 30% species with arboreal pitfall trap and 22% with the funnel trap (Table 3). However, funnel trap caught species that were not found in the arboreal pitfall trap and beating (Table 4). Fig 7. Ants numerical abundance mean (± SE) collected in different traps (numerical abundance per sample). Fig 6. Sample species richness of each trap. Species richness Simpson index Evenness Beating 27 0,89 0.34 Funnel trap 23 0,88 0.36 Arboreal pitfall 20 0,86 0.36 Table 2. Measure of diversity in the different sampling method. Sociobiology 67(4): 492-500 (December, 2020) 497 Discussion Several studies already focused on the comparison of different sampling techniques to collect arboreal ant (Kaspari, 2000; Yusah et al., 2012; Garcia-Martinez, 2018; Leponce et al., 2019). Here, we demonstrate that the “Funnel trap” can also Subfamily Espèces Arboreal pitfall (n=120) Funnel trap (n=120) Beating (n=120) Formicinae Camponotus puberulus Emery,1897 0 10 0 Camponotus compressiscapus André, 1889 0 62 0 Camponotus solon Forel, 1886 3 26 0 Camponotus acvapimensis Mayr, 1862 0 0 2 Camponutus maculatus Fabrius, 1782 2 47 0 Lepisiota sp.1 92 181 18 Oecophylla longinoda Latreille, 1802 122 1538 488 Plagiolepis alluaudi Emery, 1894 0 0 90 Plagiolepis sp.2 0 0 21 Plagiolepis sp.3 0 0 3 Polyrachis sp.1 0 1 4 Dolichoderinae Tapinoma lugubre Santschi, 1917 0 9 0 Tapinoma sp.1 0 0 2 Tapinoma sp.2 0 0 16 Myrmicinae Cataulacus traegaohdi Santschi, 1914 2 2 47 Cataulacus guineensis Smith, 1853 0 0 1 Crematogaster solenopsides Emery, 1899 163 284 1558 Crematogaster striatula Emery, 1892 1 28 42 Crematogaster africana Mayr, 1895 38 371 0 Crematogaster sp.9 23 0 1 Crematogaster nigronitens Santschi, 1917 18 1 0 Crematogaster sp.14 66 23 115 Crematogaster sp.22 0 0 25 Crematogaster sp.17 0 0 692 Crematogaster sp.21 0 0 1 Monomorium dolatu Bolton, 1987 1 0 5 Monomorium floricola Jerdon, 1851 9 97 10 Monomorium inquietum Santschi, 1926 37 57 287 Monomorium pharaonis Linnaeus, 1758 0 13 0 Monomorium sp.2 5 0 63 Monomorium sp. 3 0 0 1 Pheidole megacephala Fabricius, 1793 13 32 0 Pheidole sp.2 0 10 0 Pheidole sp.6 0 2 0 Pheidole sp.7 0 0 11 Pheidole sp.8 0 0 2 Terataner velatus Bolton, 1981 0 0 1 Tetramorium lucayanum Wheeler, 1905 1 4 0 Tetramorium quadridentatum Stitz, 1910 1 0 0 Tetramorium sp.3 4 0 0 Tetramorium sp.4 0 0 2 Ponerinae Platythyrea conradti Emery, 1899 0 1 0 Pseudomyrmicinae Tetraponera mocquerysi André, 1890 1 1 0 Total 602 2800 3670 Table 3. Similarity index (Jaccard) between ant assemblages collected by the three sampling methods. Arboreal pitfall Funnel trap Beating Arboreal pitfall 0.53 0.30 Funnel trap 0.22 Table 4. Arboreal ant species composition between three sampling method. CD Yodé, K Dosso, LMM Kouakou, K Yeo, W Dekoninck, S Konate, PK Kouassi – The “funnel trap”: a new sampling method for arboreal ants498 natural and disturbed habitats as the “funnel trap” can capture some larger ant species such as, Camponotus puberulus Emery, 1897, Camponotus compressiscapus André, 1889, Platythyrea conradti and smaller ant species like Pheidole sp.2, Monomorium floricola Jerdon, 1851, Plagiolepis alluaudi Emery, 1894 and Tapinoma lugubre Santschi, 1917. Interestingly, 48 hours after bait placement in “Funnel trap”, some ant species workers were still active and alive in the traps although the bait was totally consumed. Thus, it is possible the “funnel trap” also offers unique possibility to observe the existence of competition and interactions between ant species. For example, in some trap, we have observed a high number of killed workers of both Camponotus solon Forel, 1886 and Oecophylla longinoda Latreille, 1802, suggesting a strong competition between these two ant species in canopy. The funnel trap is an efficient sampling technique to the study of arboreal canopy ant communities. In addition, it allows to capture several other orders of insects like Blattodea, Diptera, other Hymenoptera, Coleoptera, Orthoptera and Lepidoptera. Funnel trap can also be used for sampling in the agroecosystem canopy or in other natural area besides savanna or forest. Funnel traps collect also ant species that will not easily be collected with the usually used arboreal pitfall trap. Acknowledgements The authors would like to express their sincere thanks to the African Excellence Center on Climate Change, Biodiversity and Sustainable Agriculture of Felix Houphouët Boigny University of Côte d’Ivoire for funding our research. We also thank Kouassi Dorgeles and, Koffi Kouame François for their useful help during this work. This paper is a result of several ant-course training projects with financial support from the Belgian Directorate-General for Development Cooperation (DGD), Global Taxonomy Initiative, within the framework of the CEBioS programme. Authors’ Contributions Christine Dakélé YODÉ: Design of the work, methodology, data collection, data analysis, interpretation of data for the work and writing the original draft. Wouter DEKONINCK, Kolo YEO: Contributed to revising draft, and Final approval of the version to be published. Lombart M. Maurice KOUAKOU, Kanvaly DOSSO, Souleymane KONATE, Phillipe Kouassi KOUASSI: Contributed to conceptualization, revising draft critically for important intellectual content and final approval of the version to be published. REFERENCES Abbadie, L., Gignoux J., Lepage, M., & Roux, X.L. (2006). Environmental constraints on living organisms. In Abbadie L., Gignoux J., Roux X.L. & Lepage M. (Eds.), Lamto (pp. 45-61). Springer, New York. be an efficient sampling method which requires a simple logistic, to assess arboreal ant diversity. The funnel trap” was able to catch more species from different subfamilies than the arboreal pitfall trap during a sampling campaign. On contrary, it caught less ant species than trees beating method. Probably beating also collects terricolous ant species that were encountered in the foliage. The fact that the “funnel trap” has caught more ant species than the arboreal pitfall trap might be due to the low chance of escape for ants once entering the trap. Also, this could explain the relative high rate of sampling coverage of 70% and the high number of unique species collected in comparison to the other sampling methods. The difference observed in species richness between the different sampling methods could be explained by the voracity of the ants during the dry season when there is an extra need for water and food. Indeed, baits from funnel trap and arboreal pitfall were entirely consumed or transported by the ants outside the traps. Sousa-souto et al. (2016) already mentioned that the dry season could have negative effects on the arboreal ant species richness because the loss of leaves in most tree species decreases the connectivity between tree canopies with a consequent reduction of resource availability. Sometimes traps were monopolised by a single species, leading less competitive species giving up the bait and not been collected (Garcia-Martinez, 2015). We found a high diversity for all methods, but on contrast low values for evenness. This finding matches with the study of Basset et al. (2003a) and Yusah et al. (2018) who reported that ant assemblages of tropical forest canopy are often characterized by a high diversity. Concerning the low values of evenness, a possible explanation may be that bait food attracts more species of ants that have a high recruitment rate and consequently the monopolization of baits by some dominant species (Leponce et al., 2019). These findings demonstrated that, the “funnel trap” could be therefore considered as a suitable trap to estimate the diversity of canopy ant communities and to study the structure of the canopy ant mosaic in natural and modified habitats. Beating caught more ants than the “funnel trap”, followed by arboreal pitfall trap. A plausible explanation is that generally beating was done between 2 and 3m above the ground in the foliage of trees or shrubs. At this height also, it is possible ant species that nest on the ground and occasionally forage in the trees being collected (Klimes 2017, and Vasconcelos et al., 2019). The three sampling methods generally yielded similar ant species composition. Although some studies showed that the specific composition of canopy ants varies with habitat heterogeneity (Dambros et al., 2018), here, the similarity of ant species composition observed after comparison of the three different sampling methods may be explained by the similar plant compositions of the two forest formations (Gnaoré et al., 2018). Nevertheless, the funnel trap method caught a greater number of unique species than the other methods. Therefore we would recommend it to study arboreal ants in both Sociobiology 67(4): 492-500 (December, 2020) 499 Agosti, D. & Alonso, L.E. (2000). The ALL protocol: a standard protocol for the collection of ground-dwelling ants. In Agosti, D., Majer, J., Alonso, L.E. & Schultz, T. (Eds.), Ants: standard methods for measuring and monitoring biodiversity. Smithsonian Press, Washington, pp. 204-206, doi: 10.5281/zenodo.16183. AntWeb. Available from https://www.antweb.org. (Accessed date: 12 November 2019) Basset, Y., Springate, N.D., Aberlenc, H.P. & Delvare, G. (1997). A review of methods for sampling arthropods in tree canopies. Canopy Arthropods, 35: 27-52. Basset, Y., Kitching, R.L., Miller,SE. & Novotny, V. (2003b). Arthropods of tropical forests: spatio-temporal dynamics and resource use in the canopy. Cambridge: University Press, 490 p. Bolton, B. (1980). The ant tribe Tetramoriini: The genus Tetramorium Mayr in the Ethiopian zoogeographical region. Bulletin of the British Museum (Natural History) (Entomology), 40: 193-384. Bolton, B. (1982). Afrotropical species of the myrmicine ant genera Cardiocondyla, Leptothorax, Melissotarsus, Messor, and Cataulacus. Bulletin of the British Museum (Natural History) (Entomology), 45: 307-370. Bolton, B. (1987). A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr. Bulletin of the British Museum (Natural History) (Entomology), 54: 263-452. Castaño-Meneses, G. (2014). Trophic guild structure of a canopy ant community in a mexican tropical deciduous forest. Sociobiology, 61: 35-42. doi: 10.13102/sociobiology.v61i1.35-4 Chapin, K.J. & Smith, K.H. (2019). Vertically Stratified Arthropod Diversity in a Florida Upland Hardwood Forest. Florida Entomologist, 102: 211-215. doi: 10.1653/024. 102.0134 Dambros, J., França, V.V., Delabie, J.H.C., Marques, M.I. & Battirola, L.D. (2018). Canopy Ant Assemblage (Hymenoptera: Formicidae) in Two Vegetation Formations in the Northern Brazilian Pantanal. Sociobiology, 65: 358-369. doi: 10.13102/ sociobiology. v65i3.1932 Dejean, A., McKey, D., Gibernau, M. & Belin, M. (2000). Arboreal ant mosaic in a Cameroonian rainforest (Hymenoptera: Formicidae). Sociobiology, 35: 403-423. doi: 10.5252/ zoosystema2019v41a10. Delage, B. (1970). Etude des Fourmis arboricoles de savane. Bulletin de Liaison des Chercheurs de Lamto Mars, 1970: 22-24. Delage-Darchen, B. (1971). Contribution à l’étude écologique d’une savane de Côte d’Ivoire. Les Fourmis des strates herbacée et arborée. Biologica Gabonica, 7: 461-496. Delage-Darchen, B. (1972). Une Fourmi de Côte d’Ivoire Melissotarsus titubans Delage n.sp. Insectes Sociaux, 19: 213-226. Delage-Darchen, B. (1973). Evolution de l’aile chez les Fourmis Crematogaster (Myrmicinae) d’Afrique. Insectes Sociaux, 20: 221-242. Delage-Darchen, B. (1974). Ecologie et biologie de Crematogaster impressa Emery., fourmi savanicole d’Afrique (Hymenoptera Formicidae). Insectes Sociaux, 21: 13-34. Del Toro, I., Ribbons, R.R. & Pelini, S.L. (2012). The little things that run the world revisited: a review of ant- mediated ecosystem services and disservices (Hymenoptera: Formicidae). Myrmecological News, 17: 133-146. Fisher, B.L. & Bolton, B. (2016). Ants of Africa and Madagascar: a guide to the genera. University of California Press, 251p. Floren, A., Wetzel, W. & Staab, M. (2014). The contribution of canopy species to overall ant diversity (Hymenoptera: Formicidae) in temperate and tropical ecosystems. Myrmecological News, 19: 65-74. García-Martínez, M.Á., Martínez-Tlapa, D.L., Pérez- Toledo, G.R., Quiroz-Robledo, L.N., Castaño-Meneses, G., Laborde, J. & Valenzuela-González, J.E. (2015). Taxonomic, species and functional group diversity of ants in a tropical anthropogenic landscape. Tropical Conservation Science, 8: 1017-1032. doi : 10.1177/194008291500800412 García-Martínez, M. A., Presa-Parra E., Valenzuela-González J.E. & Lasa R.. (2018). The Fruit Fly Lure CeraTrap: An Effective Tool for the Study of the Arboreal Ant Fauna (Hymenoptera: Formicidae). Journal of Insect Science, 18: 16. doi: 10.1093/jisesa/iey078 Gnahoré, E., Missa, K., Koné, M., Gueulou, N. & Bakayoko A. (2018). Dynamique et structure de la flore de la Savane Protégée des Feux dans la Réserve Scientifique de Lamto (Centre de la Côte d’Ivoire). European Scientific Journal, 14: 432. doi: 10.19044/esj.2018.v14n36p432. Hahn, D.A. & Wheeler D.E.. (2002). Seasonal Foraging Activity and Bait Preferences of Ants on Barro Colorado Island, Panama1. Biotropica, 34: 348-356, doi: 10.1111/ j.1744-7429.2002.tb00548. Hammer Ø., Harper D.A. & Ryan P.D.. (2001). PAST: paleontological statistics software package for education and data analysis. Palaeontologia electronica, 4(1), 9p. Hölldobler, B. & Wilson, E.O., (1990). The Ants. Harvard University Press, Cambridge. doi: 10.1007/978-3-662-10306-7 Klimes, P., Idigel, C., Rimandai, M., Fayle, T.M., Janda, M. Weiblen, G.D. & Novotny, V. (2012). Why are there more arboreal ant species in primary than in secondary tropical forests? Journal of Animals Ecology, 81: 1103-1112. doi: 10.1111/j.1365-2656.2012. 02002. Klimes, P. (2017). Diversity and specificity of ant-plant interactions in canopy communities: insights from primary and secondary tropical forests in New Guinea, in Oliveira, CD Yodé, K Dosso, LMM Kouakou, K Yeo, W Dekoninck, S Konate, PK Kouassi – The “funnel trap”: a new sampling method for arboreal ants500 P.S. & Koptur, S. (Eds). Ant-Plant Interactions: Impacts of Humans on Terrestrial Ecosystems, Cambridge University Press: 26-51. 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. (Eds.), “Our Planet Reviewed” 2015 large-scale biotic survey in Mitaraka, French Guiana. Zoosystema, 41: 163-179, doi: 10.5252/zoosystema2019v41a10. Lach, L., Parr, L.C. & Abbott K.L. (2010). Ant Ecology. New York: Oxford University Press Inc. 424 p. Lévieux, J. (1976). La structure du nid de quelques Fourmis arboricoles d’Afrique tropicale (Hymenoptera Formicidae). Annales de l’Université d’Abidjan, C 12: 5-22. Longino, J.T., Branstetter, M.G. & Ward, P.S. (2019). Ant diversity patterns across tropical elevation gradients: effects of sampling method and subcommunity. Ecosphere, 10. doi: 10.1002/ecs2.2798. Philpott, S.M., Greenberg, R., Bichier, P. & Perfecto, I. (2004). Impacts of major predators on tropical agroforest arthropods: comparisons within and across taxa. Oecologia, 140: 140- 149. doi 10.1007/s00442-004-1561-z Philpott, S.M. & Armbrecht I. (2006). Biodiversity in tropical agroforests and the ecological role of ants and ant diversity in predatory function. Ecological Entomology, 31: 369-377. Powell, S., Costa, A.N., Lopes, C.T. & Vasconcelos, H.L. (2011). Canopy connectivity and the availability of diverse nesting resources affect species coexistence in arboreal ants. Journal of Animal Ecology, 80: 352-360, doi: 10.1111/j.1365- 2656.2010.01779. Ribas, C.R., Schoereder, J.H., Pic, M. & Soares, S.M. (2003). Tree heterogeneity, resource availability, and larger scale processes regulating arboreal ant species richness. Austral Ecology, 28: 305-314. doi: 10.1046/j.1442-9993.2003.01290. Rigato, F. (2016). The ant genus Polyrhachis F. Smith in sub-Saharan Africa, with descriptions of ten new species. (Hymenoptera: Formicidae). Zootaxa, 4088: 1-50, doi: 10.11646/zootaxa.4088.1.1. Schonberg, L.A., Longino, J.T., Nadkarni, N.M., Yanoviak, S.P. & Gering, J.C. (2004). Arboreal Ant Species Richness in Primary Forest, Secondary Forest, and Pasture Habitats of a Tropical Montane Landscape. Biotropica, 36: 402-409. doi: 10.1646/03134 Sousa-Souto, L; Figueiredo, P.M.G; Ambrogi, B.G; Oliveira, A.C.F; Ribeiro, G.T & Neves, F.S.; (2016). Composition and Richness of Arboreal Ants in Fragments of Brazilian Caatinga: Effects of Secondary Succession. Sociobiology, 63: 762-769. doi: 10.13102/sociobiology.v63i2.909 Underwood, E.C. & Fisher B.L. (2006). The role of ants in conservation monitoring: if, when, and how. Biological conservation, 132: 166-182. doi: 10.1016/j.biocon.2006.03.022 Vasconcelos, H.L., Vilhena, J.M.S., Facure, K.G. & Albernaz, A.L.K.M. (2010). Patterns of ant species diversity and turnover across 2000 km of Amazonian floodplain forest. Journal of Biogeography, 37: 432-440. doi: 10.1111/j.1365- 2699.2009.02230. Yeo, K., Tiho, S., Ouattara, K., Konate, S., Kouakou, L.M. M. & Fofana, M. (2013). Impact de la fragmentation et de la pression humaine sur la relique forestière de l’Université d’Abobo-Adjamé (Côte d’Ivoire). Journal of Applied Biosciences, 61: 4551-4565. doi: 10.4314/jab.v61i0.85602. Yusah, K.M., Fayle, T.M., Harris, G. & Foster, W.A. (2012). Optimizing diversity assessment protocols for high canopy ants in tropical rain forest. Biotropica, 44: 73-81. doi: 10.1111/j.1744-7429. Yusah K.M., Foster, W.A., Reynolds, G. & Fayle T.M. (2018). Ant mosaics in Bornean primary rain forest high canopy depend on spatial scale, time of day, and sampling method. Peer J, 6: e4231. doi: 10.7717/peerj.4231.