Faune associated with two giant ants in northern Brazil: Dinoponera gigantea (Perty, 1833) (Ponerinae) and Paraponera clavata (Fabricius, 1775) (Paraponerinae)

Faune associated with two giant ants in northern Brazil: Dinoponera gigantea (Perty, 1833) (Ponerinae) and Paraponera clavata (Fabricius, 1775) (Paraponerinae) | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Faune associated with two giant ants in northern Brazil: Dinoponera gigantea (Perty, 1833) (Ponerinae) and Paraponera clavata (Fabricius, 1775) (Paraponerinae) Anny Kelly Castanhede Fernades Cruz, Charles Darwin Ferreira Cruz, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4908076/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract The ant nests are inhabiting by great diversity of organisms. There is few information about the nests of giant ants and their associated fauna. We study the fauna in the nests of Dinoponera gigantea (Ponerinae) and Paraponera clavata (Paraponerinae) in two localities of the state of Maranhão, Brazil. A total of 15 nests were reviewed to D. gigantea and 10 to P. clavata , recorded their associated fauna and number of chambers in each one. The total abundance of organisms recorded in nests of both species were 1833, belonging to 43 families and 30 genera/species. In the nests of D. gigantea nests were recorded 571 organisms (average ± SD = 2.48 ± 4.5 individuals by nest) while to P. clavata nests were 1,262 (2.96 ± 8.5 individuals by nest). The maximum number of chambers recorder in D. gigantea were seven, while in P. clavata were recorded 24 chambers. Insecta represent 46% of the total, Arachnida 38%, Entognatha 14%, and groups as Chilopoda, Clitellata, Diplopoda, Gastropoda and Squamata represented less than 1%. There groups as Squamata were found only in P. clavate nests. The diversity of mites and springtails was high in both species but show differences in composition. The nests or these giant ants area a very important to conservation of diversity of mani groups of myrmecophiles but also to soil fauna. Commensals Composition Myrmecophily Neotropics Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction During the construction of their nest, ants not only establish new spaces with environmental conditions stable and controlled to establish their colonies, but also create optimal and useful conditions for other organisms (Araújo et al. 2019 ). These structures, newly constructed or already established in pre-existing elements as branches or decayed fruits, mean a significant energy investment for the ants, mainly in substrate excavation and hydric and thermic regulation (Peeters et al. 1994 ; Bollazi and Rozes 2007). The success of ants in the development of their nest is so important that other organisms depend of them and used them as sites for feeding, reproduction, shelter or refuge (Sudd and Franks 1987 ; Sánchez-Piñeiro and Gómez 1995; Kronauer and Pierce 2011 ; Parmentier 2020 ). These organisms, commonly called commensals (Hölldobler and Kwapich 2022 ), can be present in different chambers of the nest, living together with ant adults and immatures, and sometimes in waste-chambers. These waste-chambers are especially important to saprophytic organisms. Some specialist ants, such as Blepharidatta conops Kempf 1967 (Diniz et al. 1998 ), feed on the cadavers of different arthropods and hunt the organisms (including ants) foraging on the waste. Many of the commensals are harmless to ants and use the variety of resources offered in the nests: the structure itself as housing, detritus when abandoned inside the nest or in an appropriate chamber (Hölldobler and Wilson 1990 ; Delabie and Jahyny 2007 ; Parker and Kronauer 2021 ). Others take advantage of the resident population and exploit it to request food from the workers, imitating, for example, the behavior of requesting food by trophollaxis, or even feeding directly on the offspring of the host ant, such as a parasite (Wilson 1971 ; Hölldobler and Wilson 1990 ; Delabie and Jahyny 2007 ; Wilson and Parker 2016 ; Hölldobler and Kwapich 2019 ; Mendonça et al. 2019 ). The species associated to the ants and their nests can invest in morphological and physiological adaptations to maintain the relationship equilibrium, as ventral suckers present on phoretic mites (Lopes et al. 2015 ; Bauman 2018 ); elongated pyriform bodies to facilitate manipulation and transport by the ants, as in several trophobiotic Hemiptera (Delabie 2001 ; Sodano et al. 2024 ); production of cuticular hydrocarbons in several groups as aphids, beetles and butterflies (Schönrogge et al. 2004 ; Endo and Itino 2013 ). There are more than 2,000 species of myrmecophilous arthropods known until now with more than 200 genera and 54 families (McIver and Stonedahl 1993 ; Rettenmeyer et al. 2011 ; Hölldobler and Kwapich 2022 ), but some estimations give the numbers of 10 000 to 100 000 species (Parker and Kronauer 2021 ). Diverse invertebrates are recorded living in or near of the nests of different ants species, and have been documented in nest of the fungus growing ants genera Apterostigma, Atta, Cyphomyrmex, Mycocepurus, Myrmicocrypta , Sericomyrmex, Trachymyrmex , as well of Paraponerinae and Ponerinae genera Mayaponera, Neoponera and Pachycondyla , and the association degree varies from mutualists, specialized simbionts to true commensals or other types of visitors (Hölldobler and Wilson 1990 ; Cushing 2012 ; Castaño-Meneses et al. 2014 , 2015 , 2017 , 2019 ; Moreira et al. 2020 ; Moleiro et al. 2021 ). Among the myrmecophilous insects, there are some Lepidoptera of the families Lycenidae and Riodinidae; Hemiptera of several families (see review by Delabie 2001 ), and Coleoptera of the families Carabidae, Coccinellidae, Paussidae, Staphylinidae (Pselaphinae), Histeridae, Scarabaeidae (Hölldobler and Wilson 1990 ; Moore and Robertson 2014 ; Parker 2016 ). Other organisms are found commonly in ant nests, such as Acari and other arachnids as scorpions, pseudoscorpions and schizomids (Cushing 2012 ; Migliorini 2019; Červená et al. 2020 ); and well as Annelida, Nematoda, Myriapoda and Mollusca (Hölldobler and Wilson 1990 ; Araújo et al. 2019 ; Dias-Soares et al. 2024 ). Most studies are focusing yet in taxonomy or simple records, but those that analyze the different types of relationships are very rare (Eickwort 1990 ; Pérez-Lachaud and Lachaud 2014 ; Ivens et al. 2016 ; Parmentier et al. 2021 ; Hölldobler and Kwapich 2022 ). The myrmecofauna of the state of Maranhão is represented by 279 species belonging to 71 genera of 10 subfamilies (Prado et al. 2019 ) with Myrmicinae as subfamily with higher species richness (126). Among these species, Dinoponera gigantea (Perty, 1833) and Paraponera clavata (Fabricius, 1775) are recognized as giant ants due to their remarkable size. These ants are recorded in the northern Amazon, Cerrado and Amazon-Cerrado transition zones, Brazil (Lenhart et al. 2013 ; Prado et al. 2019 ). Both species construct their nests mainly in the soil, in general at the base of trees, palms and other types of plants, although P. clavata has been recorded also as arboricolous (Breed and Bennett 1985 ). These species are generalist scavengers and predators, their diet includes arthropod prey, small pieces of vertebrates and other animals, nectar, part of plants as well as fecal pellets (Lenhart et al. 2013 ; Schmidt and Overal 2021 ). Finally, P. clavata has been suggested as a bioindicator species, as its foraging preferences seem related to disturbed degree in its environment (McGee and Eaton 2014 ). Our aim was to compare the fauna associated to the nests of D. gigantea and P. clavata and to carry out an analysis of the interaction networks of myrmecophiles found in the nests of these ants in the state of Maranhão, Brazil. Methodology Complete colonies of D. gigantea and P. clavata were collected in the municipalities of Caixas and Chapadinha, state of Maranhão, Brazil (Fig. 1 ). Sampling was performed during July 2019, and from January to March 2020 (Table 1 ). Sampling and translation of these ants were registered in the Biodiversity Authorization and Information System (SISBIO, Brazil: permits n o . 74298-1 and 4528-1). Table 1 Sampling localization points of Dinoponera gigantea in Caxias and Chapadinha in state of Maranhão and Paraponera clavata in the municiaplity of Caxias. UFMA = Universidade Federal do Maranhão and APA = Inhamum Protected Enviromental Area, July 2019, and January to March 2020. Municipality Locality Localization Species Nest number Caxias Ouro 4º47’S 43º20’W D. gigantea 6 Chapadinha Campus UFMA 3º44’S 43º19’W D. gigantea 9 Caxias APA 4º54’S 43º26’W P. clavata 10 Total 25 Sampling was carried out mainly in Cerrado biome, with few litter on the ground and presence of trees, babaçu palms [ Attalea speciosa (Mart. ex Spreng), Arecaceae] and shrubs layer. During sampling, carbohydrates (apple + honey) or protein (sardines in comestible oil) baits were distributed to attract the ants and localize their nest. Once the nest entrance was found, stones, sticks and other materials were removed within a 1 m radius around, from with a trench was dug 30 cm from the nest entrance. This action allowed the lateral observation of the chambers. After opening the trench, the wall was scraped with the help of a gardening shovel until it was possible to observe the beginning of the chamber. At this point, the contents of the chamber walls (about 1 cm of soil from each chamber) were carefully removed with a shovel (Fig. 2 ) and isolated in a plastic container with moistened cotton to keep alive the soil mesofauna. Associated fauna larger than 4 mm or visible to the naked eye were collected manually using entomological forceps and fixed in 70% alcohol in the field, labeled with the chamber number, nest number, date and location. To extract the associated fauna smaller than 4 mm, the chamber substrates were transported to laboratory and maintained during five days in Berlese-Tüllgren funnel (Palacios-Vargas et al. 2013 ). The fauna was sorted and counted, mites and springtails were mounted in Hoyer’s medium in order to identified them under phase contrast microscope Olympus BX51 The identification of macro and microarthropods was carried out to reach the most precise taxonomic level possible, using specialized keys (Brescovit et al. 2007 ; Sierwald 2007 ; Fujihara et al. 2011 ; Triplehorn and Johnson 2011 ) and also the review by specialist: Dr. Maria José Dias Sales (UNEB) for termites and César Augusto Galando Bernal (PPGZOO/UESC) for spiders. All invertebrates collected were labeled, mounted when necessary and deposited in the Laboratory of Social Arthropods (LABAS) at the State University of Santa Cruz (UESC), Ilhéus, Bahía, Brazil. Data analysis We analyze the similarity of fauna found in the nest chambers of D. gigantea and P. clavata according with the depth and length of the chambers, using non-metric multidimensional scaling (NMDS) analysis based on the presence-absence of species and morphospecies, with Jaccard index as distance and Monte Carlo permutations. We use the software PAST ver. 4.0 (Hammer et al.2001). The depths of the chambers were grouped into eight (to D. gigantea ) and four (to P. clavata ) categories, the stress measure was based on the Kruskal’s stress value (1964). Results A total of 25 complete nests of the two species were collected. All nests of D. gigantea (15) were located at the base of the trees or babaçu palms, while P. clavata nests (10) were located at the base of medium size shrubs (not identified species). A total of 571 organisms were found in all nests of D. gigantea nests (average ± SD = 2.48 ± 4.5 individuals by nest) while to P. clavata nests were recorded 1,262 organisms (2.96 ± 8.5 individuals by nest). The maximum number of chambers recorder in D. gigantea were seven, while in P. clavata were recorderd 24 chambers. Among the 25 studied nests of giant ants ( D. gigantea and P. clavata ), we recorded groups belonging to eight classes/subclasses, 22 orders/suborders, 43 families and 30 genera/species. Insecta represent 46% of the total, Arachnida 38%, Entognatha 14%, and groups as Chilopoda, Clitellata, Diplopoda, Gastropoda and Squamata represented less than 1% (Table 2 ). The most abundant groups in nest of D. gigantea were ants (118) follow by pseudoscorpions (81), uropodine mites of Uroobovella genus (75) and Cyphoderus springtails (61); in P. clavata were termites (360 individuals), Parasitidae mites (155), Pheidole flavens (150), Seira sp. (Collembola: 95) and Oribatida mites (92). In both ant species nests, the phylum Arthropoda was the dominant group, represented by the orders: Araneae, Sarcoptiformes (Oribatida), Mesostigmata, Trombidiformes, Opiliones, Pseudoscorpionida, Chordeumatida, Geophilomorpha, Scolopendromorpha, Diplura, Collembola (Entomobryomorpha), Blattodea, Coleoptera, Dermaptera, Hemiptera, Hymenoptera, Psocoptera, Thysanoptera, Plecoptera and Lepidoptera. In addition, a single specimen of Squamata (Amphisbaenidae), phylum Chordata, was recorded in P. clavata nest. The most abundant groups were mites (Mesostigmata, Sarcoptiformes, Tombidiformes), springtails (Entognatha), ants (Hymenoptera: Formicidae) and termites (Blattodea Isoptera). A total of 578 individuals of mites were collected, representing 31% of the total in both species of studies ants, and the most abundant were Parasitidae and Urodinychidae, this last represented firstly by the genus Uroobovella . (Table 2 ). The spiders found in the nests belongs to the genera Araneus , Attacobius , Abapeba , Parabatinga and Idiops (Table 2 ). A total of 236 individuals of springtails have been collected, representing 12% of the total abundance of organisms in both ant species nests, and the most abundant genera were Seira and Cyphoderus (Table 2 ). The Blattodea represent the 21% of the total of the individuals of the associated fauna, with Isoptera as the most abundant, representing 19% of total of associated fauna (Table 2 ). Hymenoptera were represented by 393 individuals of the Formicidae family, represent 21% of the total. The most abundant species were Roger, 1863 (277 ind.) and Strumigenys enlogata Roger 1863 (41 individuals). Pheidole flavens shown complete colonies into the nests of these giant ants species, including winged females, males and immatures. According with the NMDS graph, the depth of chambers affects the composition of fauna in both species. In the nests of D. gigantean , the deepest chambers contain the fauna of the outer chamber (Fig. 3 ), the stress recorded was 0.91. In the case of P. clavata nests, the deepest nest are in the center, and the stress was 0.18, showing a statistical suspicious significance for the similarity between fauna in the different chamber (Fig. 4 ) Discussion The ant Ph. flavens was frequent and widespread in small mixed populations of D. gigantea and P clavata and distributed in several chambers, therefore, it is believed to be a facultative resident species in nests as this ant is commonly found in leaf-litter samples. Ph. flavens is a tiny ant of approximately 1mm length, which can circulate without being inconvenienced in the nests of those giant ants. This ant possible feeds on prey remains and other resources available in detritus accumulated in different places in the chambers of these nests (Delabie et al. 2007). The presence of small species of Pheidole in the nests of Dinoponera has been sometimes reported in Dinoponera and Pheidole studies, for example, different species of Pheidole in nests of D. quadriceps by Vasconcellos et al. ( 2004 ); Pheidole rudigenis Emery, 1906 in nests of Dinoponera lucida Emery, 1901 and Pheidole dinophila Wilson 2003 in nest of Dinoponera australis [= Dinoponera grandis (Guérin-Menéville, 1838)] (Wilson 2003 ); and Ph. flavens in nests of P. clavata (Moreira et al. 2020 ). The other ant species found in the nests were Brachymyrmex heeri Forel, 1874, Carebara sp.1, Centromyrmex brachycola (Roger, 1861), Gnamptogenys moelleri (Forel, 1912), Hypoponera sp.1, Pheidole sp.2 (grupo diligens), Pseudomyrmex gracilis (Fabricius, 1804), Pseudomyrmex sp.1, Pseudoponera gilberti (Kempf, 1960), Solenopsis sp.1, Solenopsis sp.2, Strumigenys elongata Roger, 1863, Strumigenys perparva Brown, 1958, Strumigenys sp.1, were considered tourist species in this study (Belshaw and Bolton 1993 , 1994 ). Here we call “tourist” the ants whose workers occasionally forage outside or even inside the nests of other ants, but which are never resident. Mites were the most abundant group in the nests of D. gigantea and P. clavata with more than 30% of the total abundance. This group is frequently found in nests of different groups of ants such as Ponerinae and Formicinae in a range of environments (Arroyo et al. 2015 ; Lopes et al. 2015 ; Moreira et al. 2020 ). In Mexico Rocha et al. ( 2020 ) found mites in more than 98% of Neoponera villosa (Fabricius, 1804) nests. The Laelapidae family is cosmopolitan and includes mites living in a diversity of habitats and associations. These mites can live freely in the soil or associated with other arthropods, and some of them have parasitic habits, living as ectoparasites of mammals (Casanueva 1993 ). Silva et al. ( 2018 ) showed that a species of genus Cosmolaelaps Berlese, 1903 is associated with Neoponera inversa (Smith) and suggested that the mite uses the ant for phoresis. Rocha et al. ( 2020 ) observed another species of the same genus living in nests of Neoponera villosa (Fabricius, 1804). In this case, the authors considered this genus kleptoparasite, as this mite was observed in groups or alone on the ventral region of ant larvae feeding directly on it or from the food brought by the workers. In our study, no evidence of kleptoparasitism was observed. The mites of genus Uroobovella (Urodinychidae) probably use ant nests as shelters, or looking for food. The adults use ants for phoresy (Lehtinen 1987 ). Between the oribatid mites, species of Galumnidae maintain phoretic relationships with the Ponerinae N. villosa (Rocha et al. 2020 ). Regarding the springtails (Collembola) found in our study, the genera that stood out were Seira , Proisotoma and Cyphoderus. Cyphoderus is extremely common in ant nests, since it is abundant and found in the nests of several ant species (Castaño-Meneses et al. 2014 , 2015 ; Oliveira et al. 2023 ). Recently, Mota-Filho et al. ( 2021 ) found Cyphoderus innominatus Mills, 1938, in Atta sexdens nests in an Atlantic Forest-Cerrado transition area of the state of São Paulo. The occurrence of this genus is attributed to the large amount of resources available, as these organisms have a special attraction for the mycelium of the fungus cultivated by ants (Kistner, 1982 ). In their study with the army ant Eciton burchellii (Westwood, 1842) Rettenmeyer et al. ( 2011 ) observed more than 300 species associated with this ant. The authors point out that mites and springtails are part of this fauna, and they recorded phoretic mites from the families Scutacaridae and Pygmephoridae as well as other Uropodina mites living in garbage deposits. A large number of Collembola, including some Cyphoderus , were also found in the waste dumps. Given the frequency and abundance observed in nests of different ant species, some mites and springtails, along with other invertebrates, can be considered authentic myrmecophiles, benefiting from the social habits of their hosts. Glasier et al. ( 2018 ) preferred not to include mites and springtails in their analyses, as, according to these authors, there are few conclusions about the myrmecophily of these organisms, in addition to the difficulty of identifying these arthropods. However, at the light of our own observations, we consider mites and springtails as myrmecophiles as other authors (Rettenmeyer et al. 2011 ; Castaño-Meneses et al. 2015 ; Araújo et al. 2019 ; Castaño-Meneses et al. 2017 , 2019 ; Moreira et al. 2020 ; Rocha et al. 2020 ) . We observed the occurrence of Blatellidae and more frequently, Isoptera (Blattodea), in some chambers. In general, ants are considered termite predators (Tuma et al. 2020 ). Furthermore, the occurrence of termites in our samples is justified by the fact that certain ant nest were very close to the termite mounds, leading to the intersection of the ant chambers with the termite mound galleries. Elsewhere this is a commonly observed situation (see Santos et al., 2010 ). For these reasons, the observation of termites in giant ant nests requires a more detailed study. Some spiders use ant nests as shelter to actively hunt prey around (Cushing 2012 ). In some cases, they even feed on the ants themselves (Rosa, 2008 ). Other Arachnids, such as Pseudoscorpiones and Opiliones, may have a relationship of predation on the ant larvae in the nests and may also use the place as a shelter. Rocha et al. ( 2020 ) suggested that the relationship between pseudoscorpions and N. villosa was predation, since individuals of Chelodamus mexicolens Chamberlin, 1925 were clearly observed feeding on the ant larvae. The myriapods found in our study, represented by the orders Scolopendromorpha and Geophilomorpha, are organisms that penetrate larger ant nests where they seek a favorable humid microclimate. Individuals also usually build systems of galleries in the ground or under rocks and logs that give them access to a cavity where the animal hides. It is suspected too that, as they are predators of several types of invertebrates including small arthropods, ants could be a particularly interesting food resource for this group of animals (Voigländer 2011 ). Several immatures and adults of beetles were observed during our study, in particular, Staphylinidae, which are common in ant nest of many species (Parker 2016 ). These are organisms that present different levels of association, ranging from simple visitors to parasites. Some beetles enter to ant nests to feed on living or dead insects or even ant larvae; other eat detritus, nest residues and fungi that grow around (Lacau et al. 2001 ; Staniec and Zagaja 2008 ; Lapeva–Gjonova 2013). The Psocotera family Liposcelididae has been reported to occur in nests of Formica rufa or Formica pratensis (Ostrovsky and Georgiev 2020 ). Many species of this family are anthropophile and are widely distributed (Lienhard 1998 ). In general, these animals prefer places with a microclimatic stability: an average temperature of 30ºC and relative humidity of 70%, ideal conditions for the realization of their biological cycle (Rees and Walker 1990 ). The ant nests offer these conditions that must be particularly favorable for these organisms. Other groups of invertebrates were also sampled, such as Chordeumatida, Dermaptera, Diplura, Hemiptera, Opiliones and Thysanoptera. All of these have appeared with some frequency in studies that address commensals of poneromorph ants (Araújo et al. 2019 ; Moreira et al. 2020 ; Rocha et al. 2020 ). Most of them are detritivorous animals, acting mainly in decomposition and nutrient cycling (Hopkin and Read 1992 ). Their occurrence in ant nests studied seem due to its type of diet and the high amount of decomposing organic material available as a food source. On the other hand, Plecoptera appears for the first time as a myrmecophilous organism. The diet of adults is variable, with some genera depending on spores and pollen while others feed on green algae or lichens (Tierno de Figueroa and López-Rodríguez 2019 ). Its occurrence here is certainly casual, since we found a specimen only once. The record of an Amphisbaenidae (unidentified) was also due to the fact that these animals spend the entirety of their life cycle underground, and that their diet is based on the consumption of arthropods, especially ants (Esteves et al. 2008 ; Balestrin and Cappellari 2011 ), which could explain its presence in the ant nests. In the ant nest, several observations of associations between ants and reptiles have been reported, mainly as predation (Goldsbrough et al. 2006 ; Whitfield and Donnelly 2006 ; Balestrin and Cappellari 2011 ) or inquinilism (Oliveira and Della Lucia 1993 ). Gastropod have been recorded in the nests of Diacamma , Mayaponera and Neoponera (Verdcourt 2002 ; Witte et al. 2002 ; Eguchi et al. 2005 ; Araújo et al. 2019 ; Castaño-Meneses et al. 2019 ; Dias-Soares et al. 2024 ). In these nests, the differents interactions of myrmecophily and also the gastropod species were varaible, as example, in netss of Leptgenys processionalis distinguenda (Jerdon, 1851) events of facultative commensalisms or obligate symbiosis were observed only between the ant and Allopeas myrmecophilos (Janssen and Witte, 2002 ) according with Witte et al. ( 2002 ). Also Eguchi et al. ( 2005 ) recorded, in nests of Diacamma scalpratum (Smith, 1858), the trasport of four species of gastropods by ant workers and also the active entry of these mollusks into the ant’s nests. In the Neotropics, of the eight species of gastropods recorded in nests of N. verenae there are observations of interactions such as antennal touches between ant workers and six species of gastropods, in addition to the movement of mollusks in the nest chambers in the field and in the laboratory recorded by Dias-Soares et al. ( 2024 ). Other groups considered as commensals in the nests were symphylans and other groups corroborate the studies by Araújo et al. ( 2019 ); Castaño-Meneses et al. ( 2019 ); Hölldobler and Wilson ( 1990 ); Lapeva-Gjonova ( 2013 ) and Rocha et al. ( 2020 ), which include these organisms as commensals in ant nests of different species. Table 2 Fauna associated to nests of the giant ant species D. gigantea and P. clavata , collected in three municipalities in the state of Maranhão. Class/subclass Order/Suborder Family Genus/Species Abundance Arachnida Araneae Araneidae Araneus 2 Arachnida Araneae Corinnidae 6 Arachnida Araneae Corinnidae Attacobius 4 Arachnida Araneae Corinnidae Abapeba 10 Arachnida Araneae Nemesiidae 2 Arachnida Araneae Theridiidae 1 Arachnida Araneae Ctenidae Parabatinga 3 Arachnida Mesostigmata (Oribatida) 77 Arachnida Mesostigmata (Oribatida) Ameroseiidae 1 Arachnida Mesostigmata (Oribatida) Ascidae 3 Arachnida Mesostigmata (Oribatida) Chaetodactylidae 1 Arachnida Mesostigmata (Oribatida) Laelapidae 24 Arachnida Mesostigmata (Oribatida) Macrochelidae 3 Arachnida Mesostigmata (Oribatida) Parasitidae 153 Arachnida Mesostigmata (Oribatida) Phytoseiidae 6 Arachnida Mesostigmata (Oribatida) Podocinidae Podocinum 1 Arachnida Mesostigmata (Oribatida) Rhodacaridae 12 Arachnida Mesostigmata (Oribatida) Uropodidae 9 Arachnida Mesostigmata (Oribatida) Uropodidae Uropoda sp. 3 Arachnida Mesostigmata (Oribatida) Uropodidae Trachyuropoda 3 Arachnida Opiliones 2 Arachnida Pseudoscorpiones 88 Arachnida Sarcoptiformes Glycyphagidae 8 Arachnida Sarcoptiformes 99 Arachnida Sarcoptiformes Carabodidae 2 Arachnida Sarcoptiformes Euphthiracaridae 1 Arachnida Sarcoptiformes Galumnidae 2 Arachnida Sarcoptiformes Haplochthoniidae 1 Arachnida Sarcoptiformes Malaconothridae 1 Arachnida Sarcoptiformes Nothridae 2 Arachnida Sarcoptiformes Oppidae 9 Arachnida Sarcoptiformes (Astigmata) Acaridae 1 Arachnida Sarcoptiformes (Astigmata) Histiostomatidae 2 Arachnida Sarcoptiformes (Astigmata) Histiostomatidae Histiostoma sp. 2 Arachnida Sarcoptiformes (Astigmata) Hyadesiidae 1 Arachnida Sarcoptiformes (Astigmata) 1 Arachnida Tombidiformes Trombidiidae 4 Arachnida Araneae Idiopidae Idiops 1 Arachnida Mesostigmata Urodinychidae Uroobovella 146 Chilopoda Geophilomorpha 1 Chilopoda Scolopendromorpha 1 Chilopoda 6 Chilopoda 3 Clitellata 2 Diplopoda Chordeumatida 18 Entognatha Collembola/Entomobryomorpha Seira 103 Entognatha Diplura 1 Entognatha Diplura Campodeidae 15 Entognatha Diplura Parajapygidae 8 Entognatha Collembola/Entomobryomorpha Istomomidae Proisotoma 46 Entognatha Collembola/Entomobryomorpha Paronellidae Cyphoderus 66 Entognatha Collembola/Entomobryomorpha Isotomidae Folsomina 21 Gastropoda 5 Insecta Blattodae/Blattaria 20 Insecta Blattodae/Termitidae Syntermes sp. 7 Insecta Blattodae/Termitidae Subulitermes 363 Insecta Coleoptera 11 Insecta Coleoptera Staphylinidae 4 Insecta Dermaptera 2 Insecta Hemiptera Aphididae 1 Insecta Hymenoptera Formicidae 26 Insecta Hymenoptera Formicidae Brachymyrmex heeri Forel, 1874 9 Insecta Hymenoptera Formicidae Carebara sp.1 4 Insecta Hymenoptera Formicidae Centromyrmex brachycola (Roger, 1861) 1 Insecta Hymenoptera Formicidae Gnamptogenys moelleri (Forel, 1912) 1 Insecta Hymenoptera Formicidae 1 Insecta Hymenoptera Formicidae Hypoponera sp. 1 3 Insecta Hymenoptera Formicidae Pheidole flavens Roger, 1863 277 Insecta Hymenoptera Formicidae Pheidole sp.2 (grupo diligens) 2 Insecta Hymenoptera Formicidae Solenopsis sp.1 23 Insecta Hymenoptera Formicidae Solenopsis sp.2 1 Insecta Hymenoptera Formicidae Strumigenys sp.1 1 Insecta Hymenoptera Formicidae Strumigenys elongata Roger, 1863 41 Insecta Hymenoptera Formicidae Pseudomyrmex gracilis (Fabricius, 1804) 1 Insecta Hymenoptera Formicidae Strumigenys perparva Brown, 1958 1 Insecta Hymenoptera Formicidae Pseudomyrmex sp.1 1 Insecta Plecoptera 1 Insecta Psocoptera 4 Insecta Psocoptera Caeciliusidae 1 Insecta Psocoptera Liposcelididae 2 Insecta Thysanoptera Thripidae 1 Insecta Coleoptera Cantharidae 2 Insecta Coleoptera Chrysomelidae 4 Insecta Hemiptera 3 Insecta Lepidoptera (Imaturo) 16 Insecta Diptera (Imaturo) 4 Reptilia Squamata Amphisbaenidae 1 Conclusions The nest of giant ant constitute important habitat to high diversity of commensals and myrmecophilous organisms. Nevertheless there are many species share in nests of D. gigantea and P. clavata , composition of associated fauna is particular in each species, and also the spatial distribution, according with the structure complexity of the nest. Declarations Declaration Conflict of interest: The authors declare there are no conflicts of interest. Ethics approval: All experiments followed the appropriate ethical guidelines. Aknowldgements This study was supported by the Brazilian Council of Research and Scientific Development (CNPq Grant for CSFM PQ 307859/2018-5 and JHCD PQ 304629/2018-9). AKCF (CAPES 88887.485149/2020-00) and CDFC (CAPES 88887.485154/2020-00) acknowledges the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the Grant received. We thanks to Drs. Jean-Paul Lachaud and Gabriela Pérez-Lachau the invitation to participate with this work. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4908076","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":342281403,"identity":"96165ea0-88bf-43c8-8055-d3b5e86d10a8","order_by":0,"name":"Anny Kelly Castanhede Fernades Cruz","email":"","orcid":"","institution":"UESC: Universidade Estadual de Santa Cruz","correspondingAuthor":false,"prefix":"","firstName":"Anny","middleName":"Kelly Castanhede Fernades","lastName":"Cruz","suffix":""},{"id":342281404,"identity":"f1450a54-81bb-47e2-822c-d96b6113203c","order_by":1,"name":"Charles Darwin Ferreira Cruz","email":"","orcid":"","institution":"UESC: Universidade Estadual de Santa Cruz","correspondingAuthor":false,"prefix":"","firstName":"Charles","middleName":"Darwin Ferreira","lastName":"Cruz","suffix":""},{"id":342281405,"identity":"3d01b728-d419-42ae-bf3a-24770a2db707","order_by":2,"name":"Cléa S.F. Mariano","email":"","orcid":"","institution":"UESC: Universidade Estadual de Santa Cruz","correspondingAuthor":false,"prefix":"","firstName":"Cléa","middleName":"S.F.","lastName":"Mariano","suffix":""},{"id":342281406,"identity":"a1e4423c-ca10-43cb-b231-03cf0317a9d8","order_by":3,"name":"Jacques H.C. Delabie","email":"","orcid":"","institution":"CEPLAC: Comissao Executiva do Plano da Lavoura Cacaueira","correspondingAuthor":false,"prefix":"","firstName":"Jacques","middleName":"H.C.","lastName":"Delabie","suffix":""},{"id":342281407,"identity":"b338b21d-363d-4554-a027-f101ecc2fb09","order_by":4,"name":"Gabriela Castaño-Meneses","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA9ElEQVRIiWNgGAWjYFACHgjFz958DMxgYydSi4Rkz7E0BoYEoBZmYrUY3MgxA2thIKRF3v3sMYkfNXV1QC3fHnz8sU2ej5mB8cPHHNxaDM/kpQHddFhC8szb7YYzEm4btjEzMEvO3IZHS0OOmTQD2wEJvuO526R5Em4zArWwMfPi09L/BqjlX50Ew4GcZyAt9gS1yEsAbQGaLCFwIocNpCWRoBYDiXfJlr19hyVn9hwzk5yRdju5jZmxGa9f5PtzD9748a2OHxiVzyQ+2Ny2nd/efPDDR3y2HGBgkUATY2zArR5kSwMD8we8KkbBKBgFo2AUAABR9k8klVZ0NQAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0002-5405-5221","institution":"Universidad Nacional Autonoma de Mexico Facultad de Ciencias","correspondingAuthor":true,"prefix":"","firstName":"Gabriela","middleName":"","lastName":"Castaño-Meneses","suffix":""}],"badges":[],"createdAt":"2024-08-13 15:12:10","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4908076/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4908076/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":64598638,"identity":"6dda872c-2d2b-4bf6-a4a5-bdc609cd284d","added_by":"auto","created_at":"2024-09-16 11:25:05","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":303148,"visible":true,"origin":"","legend":"\u003cp\u003eLocalization of sampling points. a) \u003cem\u003eDinoponera gigantea\u003c/em\u003e sampling points in two municipalities in the state of Maranhão. (\u003cstrong\u003eB\u003c/strong\u003e) \u003cem\u003eParaponera clavata\u003c/em\u003e sampling points in the Protected Enviromental Area of Inhamum, Caxias, Maranhão.\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-4908076/v1/cc2a171faf8ea766d55692ae.png"},{"id":64598637,"identity":"66d72a4d-76cc-4906-840e-df99568c2376","added_by":"auto","created_at":"2024-09-16 11:25:05","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":94848,"visible":true,"origin":"","legend":"\u003cp\u003eSampling method of nests of \u003cem\u003eD. gigantea\u003c/em\u003e and \u003cem\u003eP. clavata\u003c/em\u003e. a) Trench around the nest before excavation (red arrow). b) Soil removal from nest chambers\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-4908076/v1/8d7a536099abcb0bcbf1b10a.png"},{"id":64598636,"identity":"b97b00cf-90e8-4278-909d-ebfd504acf47","added_by":"auto","created_at":"2024-09-16 11:25:05","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":561113,"visible":true,"origin":"","legend":"\u003cp\u003eNMDS analysis to the similarity of the associated fauna comparing different \u003cem\u003eD. gigantea\u003c/em\u003e nests\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-4908076/v1/dab3df0aa00f43386cabb1b7.png"},{"id":64598635,"identity":"96c45993-177e-4683-bcd1-216a3a968376","added_by":"auto","created_at":"2024-09-16 11:25:05","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":202409,"visible":true,"origin":"","legend":"\u003cp\u003eNMDS analysis to the similarity of the associated fauna comparing different \u003cem\u003eP. clavata\u003c/em\u003e nests\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-4908076/v1/04e5765d60ab19ffb7996351.png"},{"id":64599594,"identity":"be2a1d39-98c8-4e7e-8166-812ac056be47","added_by":"auto","created_at":"2024-09-16 11:41:08","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2154517,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4908076/v1/4c10b640-a3e6-4a11-90ef-b3c45da09cab.pdf"}],"financialInterests":"","formattedTitle":"Faune associated with two giant ants in northern Brazil: Dinoponera gigantea (Perty, 1833) (Ponerinae) and Paraponera clavata (Fabricius, 1775) (Paraponerinae)","fulltext":[{"header":"Introduction","content":"\u003cp\u003eDuring the construction of their nest, ants not only establish new spaces with environmental conditions stable and controlled to establish their colonies, but also create optimal and useful conditions for other organisms (Ara\u0026uacute;jo et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). These structures, newly constructed or already established in pre-existing elements as branches or decayed fruits, mean a significant energy investment for the ants, mainly in substrate excavation and hydric and thermic regulation (Peeters et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Bollazi and Rozes 2007). The success of ants in the development of their nest is so important that other organisms depend of them and used them as sites for feeding, reproduction, shelter or refuge (Sudd and Franks \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; S\u0026aacute;nchez-Pi\u0026ntilde;eiro and G\u0026oacute;mez 1995; Kronauer and Pierce \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Parmentier \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThese organisms, commonly called commensals (H\u0026ouml;lldobler and Kwapich \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), can be present in different chambers of the nest, living together with ant adults and immatures, and sometimes in waste-chambers. These waste-chambers are especially important to saprophytic organisms. Some specialist ants, such as \u003cem\u003eBlepharidatta conops\u003c/em\u003e Kempf 1967 (Diniz et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1998\u003c/span\u003e), feed on the cadavers of different arthropods and hunt the organisms (including ants) foraging on the waste.\u003c/p\u003e \u003cp\u003eMany of the commensals are harmless to ants and use the variety of resources offered in the nests: the structure itself as housing, detritus when abandoned inside the nest or in an appropriate chamber (H\u0026ouml;lldobler and Wilson \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Delabie and Jahyny \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Parker and Kronauer \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOthers take advantage of the resident population and exploit it to request food from the workers, imitating, for example, the behavior of requesting food by trophollaxis, or even feeding directly on the offspring of the host ant, such as a parasite (Wilson \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e1971\u003c/span\u003e; H\u0026ouml;lldobler and Wilson \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Delabie and Jahyny \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Wilson and Parker \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; H\u0026ouml;lldobler and Kwapich \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Mendon\u0026ccedil;a et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe species associated to the ants and their nests can invest in morphological and physiological adaptations to maintain the relationship equilibrium, as ventral suckers present on phoretic mites (Lopes et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Bauman \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2018\u003c/span\u003e); elongated pyriform bodies to facilitate manipulation and transport by the ants, as in several trophobiotic Hemiptera (Delabie \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Sodano et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2024\u003c/span\u003e); production of cuticular hydrocarbons in several groups as aphids, beetles and butterflies (Sch\u0026ouml;nrogge et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Endo and Itino \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThere are more than 2,000 species of myrmecophilous arthropods known until now with more than 200 genera and 54 families (McIver and Stonedahl \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Rettenmeyer et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; H\u0026ouml;lldobler and Kwapich \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), but some estimations give the numbers of 10 000 to 100 000 species (Parker and Kronauer \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDiverse invertebrates are recorded living in or near of the nests of different ants species, and have been documented in nest of the fungus growing ants genera \u003cem\u003eApterostigma, Atta, Cyphomyrmex, Mycocepurus, Myrmicocrypta\u003c/em\u003e, \u003cem\u003eSericomyrmex, Trachymyrmex\u003c/em\u003e, as well of Paraponerinae and Ponerinae genera \u003cem\u003eMayaponera, Neoponera\u003c/em\u003e and \u003cem\u003ePachycondyla\u003c/em\u003e, and the association degree varies from mutualists, specialized simbionts to true commensals or other types of visitors (H\u0026ouml;lldobler and Wilson \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Cushing \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Casta\u0026ntilde;o-Meneses et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2014\u003c/span\u003e, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2017\u003c/span\u003e, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Moreira et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Moleiro et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Among the myrmecophilous insects, there are some Lepidoptera of the families Lycenidae and Riodinidae; Hemiptera of several families (see review by Delabie \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2001\u003c/span\u003e), and Coleoptera of the families Carabidae, Coccinellidae, Paussidae, Staphylinidae (Pselaphinae), Histeridae, Scarabaeidae (H\u0026ouml;lldobler and Wilson \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Moore and Robertson \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Parker \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Other organisms are found commonly in ant nests, such as Acari and other arachnids as scorpions, pseudoscorpions and schizomids (Cushing \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Migliorini 2019; Červen\u0026aacute; et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020\u003c/span\u003e); and well as Annelida, Nematoda, Myriapoda and Mollusca (H\u0026ouml;lldobler and Wilson \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Ara\u0026uacute;jo et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Dias-Soares et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Most studies are focusing yet in taxonomy or simple records, but those that analyze the different types of relationships are very rare (Eickwort \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; P\u0026eacute;rez-Lachaud and Lachaud \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Ivens et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Parmentier et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; H\u0026ouml;lldobler and Kwapich \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe myrmecofauna of the state of Maranh\u0026atilde;o is represented by 279 species belonging to 71 genera of 10 subfamilies (Prado et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) with Myrmicinae as subfamily with higher species richness (126). Among these species, \u003cem\u003eDinoponera gigantea\u003c/em\u003e (Perty, 1833) and \u003cem\u003eParaponera clavata\u003c/em\u003e (Fabricius, 1775) are recognized as giant ants due to their remarkable size. These ants are recorded in the northern Amazon, Cerrado and Amazon-Cerrado transition zones, Brazil (Lenhart et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Prado et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBoth species construct their nests mainly in the soil, in general at the base of trees, palms and other types of plants, although \u003cem\u003eP. clavata\u003c/em\u003e has been recorded also as arboricolous (Breed and Bennett \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1985\u003c/span\u003e). These species are generalist scavengers and predators, their diet includes arthropod prey, small pieces of vertebrates and other animals, nectar, part of plants as well as fecal pellets (Lenhart et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Schmidt and Overal \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Finally, \u003cem\u003eP. clavata\u003c/em\u003e has been suggested as a bioindicator species, as its foraging preferences seem related to disturbed degree in its environment (McGee and Eaton \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOur aim was to compare the fauna associated to the nests of \u003cem\u003eD. gigantea\u003c/em\u003e and \u003cem\u003eP. clavata\u003c/em\u003e and to carry out an analysis of the interaction networks of myrmecophiles found in the nests of these ants in the state of Maranh\u0026atilde;o, Brazil.\u003c/p\u003e"},{"header":"Methodology","content":"\u003cp\u003eComplete colonies of \u003cem\u003eD. gigantea\u003c/em\u003e and \u003cem\u003eP. clavata\u003c/em\u003e were collected in the municipalities of Caixas and Chapadinha, state of Maranh\u0026atilde;o, Brazil (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Sampling was performed during July 2019, and from January to March 2020 (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Sampling and translation of these ants were registered in the Biodiversity Authorization and Information System (SISBIO, Brazil: permits n\u003csup\u003eo\u003c/sup\u003e. 74298-1 and 4528-1).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSampling localization points of \u003cem\u003eDinoponera gigantea\u003c/em\u003e in Caxias and Chapadinha in state of Maranh\u0026atilde;o and \u003cem\u003eParaponera clavata\u003c/em\u003e in the municiaplity of Caxias. UFMA\u0026thinsp;\u003cem\u003e=\u003c/em\u003e\u0026thinsp;Universidade Federal do Maranh\u0026atilde;o and APA\u0026thinsp;=\u0026thinsp;Inhamum Protected Enviromental Area, July 2019, and January to March 2020.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMunicipality\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLocality\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLocalization\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNest number\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCaxias\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eOuro\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4\u0026ordm;47\u0026rsquo;S 43\u0026ordm;20\u0026rsquo;W\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eD. gigantea\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChapadinha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCampus UFMA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3\u0026ordm;44\u0026rsquo;S 43\u0026ordm;19\u0026rsquo;W\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eD. gigantea\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCaxias\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAPA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4\u0026ordm;54\u0026rsquo;S 43\u0026ordm;26\u0026rsquo;W\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eP. clavata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eTotal\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e25\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eSampling was carried out mainly in Cerrado biome, with few litter on the ground and presence of trees, baba\u0026ccedil;u palms [\u003cem\u003eAttalea speciosa\u003c/em\u003e (Mart. ex Spreng), Arecaceae] and shrubs layer.\u003c/p\u003e \u003cp\u003eDuring sampling, carbohydrates (apple\u0026thinsp;+\u0026thinsp;honey) or protein (sardines in comestible oil) baits were distributed to attract the ants and localize their nest.\u003c/p\u003e \u003cp\u003eOnce the nest entrance was found, stones, sticks and other materials were removed within a 1 m radius around, from with a trench was dug 30 cm from the nest entrance. This action allowed the lateral observation of the chambers. After opening the trench, the wall was scraped with the help of a gardening shovel until it was possible to observe the beginning of the chamber. At this point, the contents of the chamber walls (about 1 cm of soil from each chamber) were carefully removed with a shovel (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) and isolated in a plastic container with moistened cotton to keep alive the soil mesofauna.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eAssociated fauna larger than 4 mm or visible to the naked eye were collected manually using entomological forceps and fixed in 70% alcohol in the field, labeled with the chamber number, nest number, date and location. To extract the associated fauna smaller than 4 mm, the chamber substrates were transported to laboratory and maintained during five days in Berlese-T\u0026uuml;llgren funnel (Palacios-Vargas et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe fauna was sorted and counted, mites and springtails were mounted in Hoyer\u0026rsquo;s medium in order to identified them under phase contrast microscope Olympus BX51\u003c/p\u003e \u003cp\u003eThe identification of macro and microarthropods was carried out to reach the most precise taxonomic level possible, using specialized keys (Brescovit et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Sierwald \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Fujihara et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Triplehorn and Johnson \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) and also the review by specialist: Dr. Maria Jos\u0026eacute; Dias Sales (UNEB) for termites and C\u0026eacute;sar Augusto Galando Bernal (PPGZOO/UESC) for spiders.\u003c/p\u003e \u003cp\u003eAll invertebrates collected were labeled, mounted when necessary and deposited in the Laboratory of Social Arthropods (LABAS) at the State University of Santa Cruz (UESC), Ilh\u0026eacute;us, Bah\u0026iacute;a, Brazil.\u003c/p\u003e \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eData analysis\u003c/h2\u003e \u003cp\u003eWe analyze the similarity of fauna found in the nest chambers of \u003cem\u003eD. gigantea\u003c/em\u003e and \u003cem\u003eP. clavata\u003c/em\u003e according with the depth and length of the chambers, using non-metric multidimensional scaling (NMDS) analysis based on the presence-absence of species and morphospecies, with Jaccard index as distance and Monte Carlo permutations. We use the software PAST ver. 4.0 (Hammer et al.2001). The depths of the chambers were grouped into eight (to \u003cem\u003eD. gigantea\u003c/em\u003e) and four (to \u003cem\u003eP. clavata\u003c/em\u003e) categories, the stress measure was based on the Kruskal\u0026rsquo;s stress value (1964).\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eA total of 25 complete nests of the two species were collected. All nests of \u003cem\u003eD. gigantea\u003c/em\u003e (15) were located at the base of the trees or baba\u0026ccedil;u palms, while \u003cem\u003eP. clavata\u003c/em\u003e nests (10) were located at the base of medium size shrubs (not identified species). A total of 571 organisms were found in all nests of \u003cem\u003eD. gigantea\u003c/em\u003e nests (average\u0026thinsp;\u0026plusmn;\u0026thinsp;SD\u0026thinsp;=\u0026thinsp;2.48\u0026thinsp;\u0026plusmn;\u0026thinsp;4.5 individuals by nest) while to \u003cem\u003eP. clavata\u003c/em\u003e nests were recorded 1,262 organisms (2.96\u0026thinsp;\u0026plusmn;\u0026thinsp;8.5 individuals by nest). The maximum number of chambers recorder in \u003cem\u003eD. gigantea\u003c/em\u003e were seven, while in \u003cem\u003eP. clavata\u003c/em\u003e were recorderd 24 chambers.\u003c/p\u003e \u003cp\u003eAmong the 25 studied nests of giant ants (\u003cem\u003eD. gigantea\u003c/em\u003e and \u003cem\u003eP. clavata\u003c/em\u003e), we recorded groups belonging to eight classes/subclasses, 22 orders/suborders, 43 families and 30 genera/species. Insecta represent 46% of the total, Arachnida 38%, Entognatha 14%, and groups as Chilopoda, Clitellata, Diplopoda, Gastropoda and Squamata represented less than 1% (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe most abundant groups in nest of \u003cem\u003eD. gigantea\u003c/em\u003e were ants (118) follow by pseudoscorpions (81), uropodine mites of \u003cem\u003eUroobovella\u003c/em\u003e genus (75) and \u003cem\u003eCyphoderus\u003c/em\u003e springtails (61); in \u003cem\u003eP. clavata\u003c/em\u003e were termites (360 individuals), Parasitidae mites (155), \u003cem\u003ePheidole flavens\u003c/em\u003e (150), \u003cem\u003eSeira\u003c/em\u003e sp. (Collembola: 95) and Oribatida mites (92).\u003c/p\u003e \u003cp\u003eIn both ant species nests, the phylum Arthropoda was the dominant group, represented by the orders: Araneae, Sarcoptiformes (Oribatida), Mesostigmata, Trombidiformes, Opiliones, Pseudoscorpionida, Chordeumatida, Geophilomorpha, Scolopendromorpha, Diplura, Collembola (Entomobryomorpha), Blattodea, Coleoptera, Dermaptera, Hemiptera, Hymenoptera, Psocoptera, Thysanoptera, Plecoptera and Lepidoptera. In addition, a single specimen of Squamata (Amphisbaenidae), phylum Chordata, was recorded in \u003cem\u003eP. clavata\u003c/em\u003e nest.\u003c/p\u003e \u003cp\u003eThe most abundant groups were mites (Mesostigmata, Sarcoptiformes, Tombidiformes), springtails (Entognatha), ants (Hymenoptera: Formicidae) and termites (Blattodea Isoptera).\u003c/p\u003e \u003cp\u003eA total of 578 individuals of mites were collected, representing 31% of the total in both species of studies ants, and the most abundant were Parasitidae and Urodinychidae, this last represented firstly by the genus \u003cem\u003eUroobovella\u003c/em\u003e. (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe spiders found in the nests belongs to the genera \u003cem\u003eAraneus\u003c/em\u003e, \u003cem\u003eAttacobius\u003c/em\u003e, \u003cem\u003eAbapeba\u003c/em\u003e, \u003cem\u003eParabatinga\u003c/em\u003e and \u003cem\u003eIdiops\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eA total of 236 individuals of springtails have been collected, representing 12% of the total abundance of organisms in both ant species nests, and the most abundant genera were \u003cem\u003eSeira\u003c/em\u003e and \u003cem\u003eCyphoderus\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe Blattodea represent the 21% of the total of the individuals of the associated fauna, with Isoptera as the most abundant, representing 19% of total of associated fauna (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eHymenoptera were represented by 393 individuals of the Formicidae family, represent 21% of the total. The most abundant species were Roger, 1863 (277 ind.) and \u003cem\u003eStrumigenys enlogata\u003c/em\u003e Roger 1863 (41 individuals). \u003cem\u003ePheidole flavens\u003c/em\u003e shown complete colonies into the nests of these giant ants species, including winged females, males and immatures.\u003c/p\u003e \u003cp\u003eAccording with the NMDS graph, the depth of chambers affects the composition of fauna in both species. In the nests of \u003cem\u003eD. gigantean\u003c/em\u003e, the deepest chambers contain the fauna of the outer chamber (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e), the stress recorded was 0.91. In the case of \u003cem\u003eP. clavata\u003c/em\u003e nests, the deepest nest are in the center, and the stress was 0.18, showing a statistical suspicious significance for the similarity between fauna in the different chamber (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e)\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe ant \u003cem\u003ePh. flavens\u003c/em\u003e was frequent and widespread in small mixed populations of \u003cem\u003eD. gigantea\u003c/em\u003e and \u003cem\u003eP clavata\u003c/em\u003e and distributed in several chambers, therefore, it is believed to be a facultative resident species in nests as this ant is commonly found in leaf-litter samples. \u003cem\u003ePh. flavens\u003c/em\u003e is a tiny ant of approximately 1mm length, which can circulate without being inconvenienced in the nests of those giant ants. This ant possible feeds on prey remains and other resources available in detritus accumulated in different places in the chambers of these nests (Delabie et al. 2007).\u003c/p\u003e \u003cp\u003eThe presence of small species of \u003cem\u003ePheidole\u003c/em\u003e in the nests of \u003cem\u003eDinoponera\u003c/em\u003e has been sometimes reported in \u003cem\u003eDinoponera\u003c/em\u003e and \u003cem\u003ePheidole\u003c/em\u003e studies, for example, different species of \u003cem\u003ePheidole\u003c/em\u003e in nests of \u003cem\u003eD. quadriceps\u003c/em\u003e by Vasconcellos et al. (\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2004\u003c/span\u003e); \u003cem\u003ePheidole rudigenis\u003c/em\u003e Emery, 1906 in nests of \u003cem\u003eDinoponera lucida\u003c/em\u003e Emery, 1901 and \u003cem\u003ePheidole dinophila\u003c/em\u003e Wilson \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2003\u003c/span\u003e in nest of \u003cem\u003eDinoponera australis\u003c/em\u003e [=\u0026thinsp;\u003cem\u003eDinoponera grandis\u003c/em\u003e (Gu\u0026eacute;rin-Men\u0026eacute;ville, 1838)] (Wilson \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2003\u003c/span\u003e); and \u003cem\u003ePh. flavens\u003c/em\u003e in nests of \u003cem\u003eP. clavata\u003c/em\u003e (Moreira et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe other ant species found in the nests were \u003cem\u003eBrachymyrmex heeri\u003c/em\u003e Forel, 1874, \u003cem\u003eCarebara\u003c/em\u003e sp.1, \u003cem\u003eCentromyrmex brachycola\u003c/em\u003e (Roger, 1861), \u003cem\u003eGnamptogenys moelleri\u003c/em\u003e (Forel, 1912), \u003cem\u003eHypoponera\u003c/em\u003e sp.1, \u003cem\u003ePheidole\u003c/em\u003e sp.2 (grupo diligens), \u003cem\u003ePseudomyrmex gracilis\u003c/em\u003e (Fabricius, 1804), \u003cem\u003ePseudomyrmex\u003c/em\u003esp.1, \u003cem\u003ePseudoponera gilberti\u003c/em\u003e (Kempf, 1960), \u003cem\u003eSolenopsis\u003c/em\u003e sp.1, \u003cem\u003eSolenopsis\u003c/em\u003e sp.2, \u003cem\u003eStrumigenys elongata\u003c/em\u003e Roger, 1863, \u003cem\u003eStrumigenys perparva\u003c/em\u003e Brown, 1958, \u003cem\u003eStrumigenys\u003c/em\u003e sp.1, were considered tourist species in this study (Belshaw and Bolton \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1993\u003c/span\u003e, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1994\u003c/span\u003e). Here we call \u0026ldquo;tourist\u0026rdquo; the ants whose workers occasionally forage outside or even inside the nests of other ants, but which are never resident.\u003c/p\u003e \u003cp\u003eMites were the most abundant group in the nests of \u003cem\u003eD. gigantea\u003c/em\u003e and \u003cem\u003eP. clavata\u003c/em\u003e with more than 30% of the total abundance. This group is frequently found in nests of different groups of ants such as Ponerinae and Formicinae in a range of environments (Arroyo et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Lopes et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Moreira et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In Mexico Rocha et al. (\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) found mites in more than 98% of \u003cem\u003eNeoponera villosa\u003c/em\u003e (Fabricius, 1804) nests.\u003c/p\u003e \u003cp\u003eThe Laelapidae family is cosmopolitan and includes mites living in a diversity of habitats and associations. These mites can live freely in the soil or associated with other arthropods, and some of them have parasitic habits, living as ectoparasites of mammals (Casanueva \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1993\u003c/span\u003e). Silva et al. (\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) showed that a species of genus \u003cem\u003eCosmolaelaps\u003c/em\u003e Berlese, 1903 is associated with \u003cem\u003eNeoponera inversa\u003c/em\u003e (Smith) and suggested that the mite uses the ant for phoresis. Rocha et al. (\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) observed another species of the same genus living in nests of \u003cem\u003eNeoponera villosa\u003c/em\u003e (Fabricius, 1804). In this case, the authors considered this genus kleptoparasite, as this mite was observed in groups or alone on the ventral region of ant larvae feeding directly on it or from the food brought by the workers. In our study, no evidence of kleptoparasitism was observed.\u003c/p\u003e \u003cp\u003eThe mites of genus \u003cem\u003eUroobovella\u003c/em\u003e (Urodinychidae) probably use ant nests as shelters, or looking for food. The adults use ants for phoresy (Lehtinen \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1987\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBetween the oribatid mites, species of Galumnidae maintain phoretic relationships with the Ponerinae \u003cem\u003eN. villosa\u003c/em\u003e (Rocha et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eRegarding the springtails (Collembola) found in our study, the genera that stood out were \u003cem\u003eSeira\u003c/em\u003e, \u003cem\u003eProisotoma\u003c/em\u003e and \u003cem\u003eCyphoderus. Cyphoderus\u003c/em\u003e is extremely common in ant nests, since it is abundant and found in the nests of several ant species (Casta\u0026ntilde;o-Meneses et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2014\u003c/span\u003e, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Oliveira et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Recently, Mota-Filho et al. (\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) found \u003cem\u003eCyphoderus innominatus\u003c/em\u003e Mills, 1938, in \u003cem\u003eAtta sexdens\u003c/em\u003e nests in an Atlantic Forest-Cerrado transition area of the state of S\u0026atilde;o Paulo. The occurrence of this genus is attributed to the large amount of resources available, as these organisms have a special attraction for the mycelium of the fungus cultivated by ants (Kistner, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1982\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn their study with the army ant \u003cem\u003eEciton burchellii\u003c/em\u003e (Westwood, 1842) Rettenmeyer et al. (\u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) observed more than 300 species associated with this ant. The authors point out that mites and springtails are part of this fauna, and they recorded phoretic mites from the families Scutacaridae and Pygmephoridae as well as other Uropodina mites living in garbage deposits. A large number of Collembola, including some \u003cem\u003eCyphoderus\u003c/em\u003e, were also found in the waste dumps. Given the frequency and abundance observed in nests of different ant species, some mites and springtails, along with other invertebrates, can be considered authentic myrmecophiles, benefiting from the social habits of their hosts.\u003c/p\u003e \u003cp\u003eGlasier et al. (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) preferred not to include mites and springtails in their analyses, as, according to these authors, there are few conclusions about the myrmecophily of these organisms, in addition to the difficulty of identifying these arthropods. However, at the light of our own observations, we consider mites and springtails as myrmecophiles as other authors (Rettenmeyer et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Casta\u0026ntilde;o-Meneses et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Ara\u0026uacute;jo et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Casta\u0026ntilde;o-Meneses et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2017\u003c/span\u003e, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Moreira et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Rocha et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e ) .\u003c/p\u003e \u003cp\u003eWe observed the occurrence of Blatellidae and more frequently, Isoptera (Blattodea), in some chambers. In general, ants are considered termite predators (Tuma et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Furthermore, the occurrence of termites in our samples is justified by the fact that certain ant nest were very close to the termite mounds, leading to the intersection of the ant chambers with the termite mound galleries. Elsewhere this is a commonly observed situation (see Santos et al., \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). For these reasons, the observation of termites in giant ant nests requires a more detailed study.\u003c/p\u003e \u003cp\u003eSome spiders use ant nests as shelter to actively hunt prey around (Cushing \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). In some cases, they even feed on the ants themselves (Rosa, \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2008\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOther Arachnids, such as Pseudoscorpiones and Opiliones, may have a relationship of predation on the ant larvae in the nests and may also use the place as a shelter. Rocha et al. (\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) suggested that the relationship between pseudoscorpions and \u003cem\u003eN. villosa\u003c/em\u003e was predation, since individuals of \u003cem\u003eChelodamus mexicolens\u003c/em\u003e Chamberlin, 1925 were clearly observed feeding on the ant larvae.\u003c/p\u003e \u003cp\u003eThe myriapods found in our study, represented by the orders Scolopendromorpha and Geophilomorpha, are organisms that penetrate larger ant nests where they seek a favorable humid microclimate. Individuals also usually build systems of galleries in the ground or under rocks and logs that give them access to a cavity where the animal hides. It is suspected too that, as they are predators of several types of invertebrates including small arthropods, ants could be a particularly interesting food resource for this group of animals (Voigl\u0026auml;nder \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSeveral immatures and adults of beetles were observed during our study, in particular, Staphylinidae, which are common in ant nest of many species (Parker \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). These are organisms that present different levels of association, ranging from simple visitors to parasites. Some beetles enter to ant nests to feed on living or dead insects or even ant larvae; other eat detritus, nest residues and fungi that grow around (Lacau et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Staniec and Zagaja \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Lapeva\u0026ndash;Gjonova 2013).\u003c/p\u003e \u003cp\u003eThe Psocotera family Liposcelididae has been reported to occur in nests of \u003cem\u003eFormica rufa\u003c/em\u003e or \u003cem\u003eFormica pratensis\u003c/em\u003e (Ostrovsky and Georgiev \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Many species of this family are anthropophile and are widely distributed (Lienhard \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). In general, these animals prefer places with a microclimatic stability: an average temperature of 30\u0026ordm;C and relative humidity of 70%, ideal conditions for the realization of their biological cycle (Rees and Walker \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e1990\u003c/span\u003e). The ant nests offer these conditions that must be particularly favorable for these organisms.\u003c/p\u003e \u003cp\u003eOther groups of invertebrates were also sampled, such as Chordeumatida, Dermaptera, Diplura, Hemiptera, Opiliones and Thysanoptera. All of these have appeared with some frequency in studies that address commensals of poneromorph ants (Ara\u0026uacute;jo et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Moreira et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Rocha et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Most of them are detritivorous animals, acting mainly in decomposition and nutrient cycling (Hopkin and Read \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1992\u003c/span\u003e). Their occurrence in ant nests studied seem due to its type of diet and the high amount of decomposing organic material available as a food source.\u003c/p\u003e \u003cp\u003eOn the other hand, Plecoptera appears for the first time as a myrmecophilous organism. The diet of adults is variable, with some genera depending on spores and pollen while others feed on green algae or lichens (Tierno de Figueroa and L\u0026oacute;pez-Rodr\u0026iacute;guez \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Its occurrence here is certainly casual, since we found a specimen only once.\u003c/p\u003e \u003cp\u003eThe record of an Amphisbaenidae (unidentified) was also due to the fact that these animals spend the entirety of their life cycle underground, and that their diet is based on the consumption of arthropods, especially ants (Esteves et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Balestrin and Cappellari \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), which could explain its presence in the ant nests. In the ant nest, several observations of associations between ants and reptiles have been reported, mainly as predation (Goldsbrough et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Whitfield and Donnelly \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Balestrin and Cappellari \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) or inquinilism (Oliveira and Della Lucia \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e1993\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eGastropod have been recorded in the nests of \u003cem\u003eDiacamma\u003c/em\u003e, \u003cem\u003eMayaponera\u003c/em\u003e and \u003cem\u003eNeoponera\u003c/em\u003e (Verdcourt \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Witte et al. \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Eguchi et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Ara\u0026uacute;jo et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Casta\u0026ntilde;o-Meneses et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Dias-Soares et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In these nests, the differents interactions of myrmecophily and also the gastropod species were varaible, as example, in netss of \u003cem\u003eLeptgenys processionalis distinguenda\u003c/em\u003e (Jerdon, 1851) events of facultative commensalisms or obligate symbiosis were observed only between the ant and \u003cem\u003eAllopeas myrmecophilos\u003c/em\u003e (Janssen and Witte, \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2002\u003c/span\u003e) according with Witte et al. (\u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Also Eguchi et al. (\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2005\u003c/span\u003e) recorded, in nests of \u003cem\u003eDiacamma scalpratum\u003c/em\u003e (Smith, 1858), the trasport of four species of gastropods by ant workers and also the active entry of these mollusks into the ant\u0026rsquo;s nests. In the Neotropics, of the eight species of gastropods recorded in nests of \u003cem\u003eN. verenae\u003c/em\u003e there are observations of interactions such as antennal touches between ant workers and six species of gastropods, in addition to the movement of mollusks in the nest chambers in the field and in the laboratory recorded by Dias-Soares et al. (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOther groups considered as commensals in the nests were symphylans and other groups corroborate the studies by Ara\u0026uacute;jo et al. (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2019\u003c/span\u003e); Casta\u0026ntilde;o-Meneses et al. (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2019\u003c/span\u003e); H\u0026ouml;lldobler and Wilson (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1990\u003c/span\u003e); Lapeva-Gjonova (\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and Rocha et al. (\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), which include these organisms as commensals in ant nests of different species.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eFauna associated to nests of the giant ant species \u003cem\u003eD. gigantea\u003c/em\u003e and \u003cem\u003eP. clavata\u003c/em\u003e, collected in three municipalities in the state of Maranh\u0026atilde;o.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eClass/subclass\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eOrder/Suborder\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFamily\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGenus/Species\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAbundance\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAraneae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAraneidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eAraneus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAraneae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCorinnidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAraneae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCorinnidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eAttacobius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAraneae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCorinnidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eAbapeba\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAraneae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNemesiidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAraneae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTheridiidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAraneae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCtenidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eParabatinga\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e77\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAmeroseiidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAscidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eChaetodactylidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLaelapidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e24\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMacrochelidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eParasitidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e153\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePhytoseiidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePodocinidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003ePodocinum\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eRhodacaridae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUropodidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUropodidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eUropoda\u003c/em\u003e sp.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata (Oribatida)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUropodidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eTrachyuropoda\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eOpiliones\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePseudoscorpiones\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e88\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eGlycyphagidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e99\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCarabodidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEuphthiracaridae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eGalumnidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHaplochthoniidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMalaconothridae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNothridae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eOppidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes (Astigmata)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAcaridae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes (Astigmata)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHistiostomatidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes (Astigmata)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHistiostomatidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eHistiostoma\u003c/em\u003e sp.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes (Astigmata)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHyadesiidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSarcoptiformes (Astigmata)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTombidiformes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTrombidiidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAraneae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIdiopidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eIdiops\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArachnida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMesostigmata\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eUrodinychidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eUroobovella\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e146\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChilopoda\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGeophilomorpha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChilopoda\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eScolopendromorpha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChilopoda\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChilopoda\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eClitellata\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDiplopoda\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eChordeumatida\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEntognatha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCollembola/Entomobryomorpha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eSeira\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e103\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEntognatha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDiplura\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEntognatha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDiplura\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCampodeidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEntognatha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDiplura\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eParajapygidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEntognatha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCollembola/Entomobryomorpha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIstomomidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eProisotoma\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e46\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEntognatha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCollembola/Entomobryomorpha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eParonellidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eCyphoderus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e66\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEntognatha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCollembola/Entomobryomorpha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIsotomidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eFolsomina\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e21\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGastropoda\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlattodae/Blattaria\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlattodae/Termitidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eSyntermes\u003c/em\u003e sp.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlattodae/Termitidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eSubulitermes\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e363\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eColeoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eColeoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eStaphylinidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDermaptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHemiptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAphididae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e26\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eBrachymyrmex heeri\u003c/em\u003e Forel, 1874\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eCarebara\u003c/em\u003e sp.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eCentromyrmex brachycola\u003c/em\u003e (Roger, 1861)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eGnamptogenys moelleri\u003c/em\u003e (Forel, 1912)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eHypoponera\u003c/em\u003e sp. 1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003ePheidole flavens\u003c/em\u003e Roger, 1863\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e277\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003ePheidole\u003c/em\u003e sp.2 (grupo diligens)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eSolenopsis\u003c/em\u003e sp.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e23\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eSolenopsis\u003c/em\u003e sp.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eStrumigenys\u003c/em\u003e sp.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eStrumigenys elongata Roger, 1863\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e41\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003ePseudomyrmex gracilis\u003c/em\u003e (Fabricius, 1804)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eStrumigenys perparva\u003c/em\u003e Brown, 1958\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003ePseudomyrmex\u003c/em\u003e sp.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePlecoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePsocoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePsocoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCaeciliusidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePsocoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLiposcelididae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eThysanoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eThripidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eColeoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCantharidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eColeoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eChrysomelidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHemiptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLepidoptera (Imaturo)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eInsecta\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDiptera (Imaturo)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eReptilia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSquamata\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAmphisbaenidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eThe nest of giant ant constitute important habitat to high diversity of commensals and myrmecophilous organisms. Nevertheless there are many species share in nests of \u003cem\u003eD. gigantea\u003c/em\u003e and \u003cem\u003eP. clavata\u003c/em\u003e, composition of associated fauna is particular in each species, and also the spatial distribution, according with the structure complexity of the nest.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eDeclaration Conflict of interest: The authors declare there are no conflicts of interest.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eEthics approval: \u0026nbsp;All experiments followed the appropriate ethical guidelines.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eAknowldgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis study was supported by the Brazilian Council of Research and Scientific Development (CNPq Grant for CSFM PQ 307859/2018-5 and JHCD PQ 304629/2018-9). AKCF (CAPES 88887.485149/2020-00) and CDFC (CAPES 88887.485154/2020-00) acknowledges the Coordena\u0026ccedil;\u0026atilde;o de Aperfei\u0026ccedil;oamento de Pessoal de N\u0026iacute;vel Superior (CAPES) for the Grant received. We thanks to Drs. Jean-Paul Lachaud and Gabriela P\u0026eacute;rez-Lachau the invitation to participate with this work.\u0026nbsp;\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAra\u0026uacute;jo ES, Koch EBA, Delabie JHC, Zeppelini D, DaRocha WD, Casta\u0026ntilde;o-Meneses G, Marinao CSF (2019) Diversity of commensals within nest of ants of the genus \u003cem\u003eNeoponera\u003c/em\u003e (Hymenoptera: Formicidae: Ponerinae) in Bahia, Brazil. Ann Soc Entomol Fr 55:291-299. https://doi.org/10.1080/00379271.2019.1629837\u003c/li\u003e\n\u003cli\u003eArroyo J, O\u0026rsquo;Grady A. 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There is few information about the nests of giant ants and their associated fauna. We study the fauna in the nests of \u003cem\u003eDinoponera gigantea\u003c/em\u003e (Ponerinae) and \u003cem\u003eParaponera clavata\u003c/em\u003e (Paraponerinae) in two localities of the state of Maranh\u0026atilde;o, Brazil. A total of 15 nests were reviewed to \u003cem\u003eD. gigantea\u003c/em\u003e and 10 to \u003cem\u003eP. clavata\u003c/em\u003e, recorded their associated fauna and number of chambers in each one. The total abundance of organisms recorded in nests of both species were 1833, belonging to 43 families and 30 genera/species. In the nests of \u003cem\u003eD. gigantea\u003c/em\u003e nests were recorded 571 organisms (average\u0026thinsp;\u0026plusmn;\u0026thinsp;SD\u0026thinsp;=\u0026thinsp;2.48\u0026thinsp;\u0026plusmn;\u0026thinsp;4.5 individuals by nest) while to \u003cem\u003eP. clavata\u003c/em\u003e nests were 1,262 (2.96\u0026thinsp;\u0026plusmn;\u0026thinsp;8.5 individuals by nest). The maximum number of chambers recorder in \u003cem\u003eD. gigantea\u003c/em\u003e were seven, while in \u003cem\u003eP. clavata\u003c/em\u003e were recorded 24 chambers. Insecta represent 46% of the total, Arachnida 38%, Entognatha 14%, and groups as Chilopoda, Clitellata, Diplopoda, Gastropoda and Squamata represented less than 1%. There groups as Squamata were found only in P. clavate nests. The diversity of mites and springtails was high in both species but show differences in composition. The nests or these giant ants area a very important to conservation of diversity of mani groups of myrmecophiles but also to soil fauna.\u003c/p\u003e","manuscriptTitle":"Faune associated with two giant ants in northern Brazil: Dinoponera gigantea (Perty, 1833) (Ponerinae) and Paraponera clavata (Fabricius, 1775) (Paraponerinae)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-09-16 11:25:00","doi":"10.21203/rs.3.rs-4908076/v1","editorialEvents":[{"type":"communityComments","content":1}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"059d6e51-1d14-428f-9f88-1d86949ca134","owner":[],"postedDate":"September 16th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2024-09-16T11:25:03+00:00","versionOfRecord":[],"versionCreatedAt":"2024-09-16 11:25:00","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-4908076","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4908076","identity":"rs-4908076","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}