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

doi:10.21203/rs.3.rs-4908076/v1

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

https://doi.org/10.21203/rs.3.rs-4908076/v1

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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

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.

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).

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)

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), Pseudomyrmexsp.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

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.

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.

Araújo ES, Koch EBA, Delabie JHC, Zeppelini D, DaRocha WD, Castaño-Meneses G, Marinao CSF (2019) Diversity of commensals within nest of ants of the genus Neoponera (Hymenoptera: Formicidae: Ponerinae) in Bahia, Brazil. Ann Soc Entomol Fr 55:291-299. https://doi.org/10.1080/00379271.2019.1629837
Arroyo J, O’Grady A. Vance H, Bolger T (2015) The mite (Acari: Oribatida, Mesostigmata) assemblages associated with Lasius flavus (Hymenoptera: Formicidae) nests and surrounding soil in an Irish grassland. Biol Environ 115(1):17-28. https://doi.org/10.3318/bioe.2015.03
Balestrin RL, Cappellari L (2011) Reproduction and feeding ecology of Amphisbaena munoai and Anops kingi (Amphisbaenia, Amphisbaenidae) in the Escudo Sul-Rio-Grandense, souther Brazil. Iheringia, Sér Zool 101(1-2):93-102. https://doi.org/10.1590/S0073-47211011000100013
Bauman J (2018) Tiny mites on a great journey – a review on scutacrarid mites as phoronts and inquilines (Heterostigmatina, Pygmephoroidea, Scutacaridae) Acarologia 58(1):192-251. https://doi.org/10.24349/acarologia/20184238
Belshaw R, Bolton B (1993) The effect of forest disturbance on the leaf litter ant fauna in Ghana. Biol Conserv 2:656-666. https://doi.org/10.1007/BF00051965
Belshaw R, Bolton B (1994) A survey of the leaf litter ant fauna in Ghana, West Africa (Hymenoptera: Formicidae). J Hym Res 3:5-16. https://biostor.org/reference/80267
Bollazzi M, Roces F (2007) To build or not to build: circulating dry air organizes collective building for climate control in the leaf-cutting ant Acromyrmex ambiguus. Anim Behav 74(5):1349-1355. https://doi.org/10.1016/j.anbehav.2007.02.021
Breed MD, Bennett B (1985) Mass recruitment to nectar sources in Paraponera clavata: a field study. Ins Soc 32:198-208. https://doi.org/10.1007/BF02224233
Brescovit AD, Rheims, CA, Bonaldo AB (2007) Araneomorpha: chave de identificação para famílias de aranhas brasileiras. Instituto Butantan, São Paulo
Casanueva ME (1993) Phylogenetic studies of the free-living and arthropod associated Laelapidae (Acari: Mesostigmata). Gayana Zool 57:21-46
Castaño-Meneses G, Palacios-Vargas JG, Delabie JHC, Santos RJ, Mariano CSF (2014) Springtails (Collembola) from nests of Ponerinae (Hymenoptera: Formicidae) ants in Brazilian cacao plantations. Flo Entomol 97(4):1862-1864. https://doi.org/10.1653/024.097.0468
Castaño-Meneses G, Palacios-Vargas, JG, Carmo AFR (2015) Colêmbolos e outros inquilinos de formigueiros de poneromorfas. In: Delabie JCH, Feitosa RM, Serrão JE, Mariano CSF, Majer JD (eds) As formigas Poneromorfas do Brasil, Editus, Ilhéus, pp 163–179
Castaño-Meneses G, Palacios-Vargas JG, Delabie JHC, Zeppelini D, Mariano CSF (2017) Springtails (Collembola) associated with nests of fungus-growing ants (Formicidae: Myrmicinae: Attini) in southern Bahia, Brazil. Flo Entomol 100(4):740-742. https://doi.org/10.1653/024.100.0421.
Castaño-Meneses G, Santos RJ, Santos JRM, Delabie JHC, Lopes LL, Mariano CSF (2019) Invertebrates associated to Ponerine ants nests in two cocoa farming systems in the southeast of the state of Bahia, Brazil. Trop Ecol 60(1):52-61. https://doi.org/10.1007/s42965-019-00006-3
Cushing PE (2012) Spider-ant associations: an update review of myrmecomorphy, myrmecophily, and myrmecophagy in spiders. Psyche 2012:151989. https://doil.org/10.1155/2012/151989
Červená M, Krajčovičová K & Christophoryová J (2020) Pseudoscorpions (Arachnida: Pseudoscorpiones) in the nests of Formica ants in Slovakia. Klapalekiana 56:205-212.
Delabie JHC (2001) Trophobiosis between Formicidae and Hemiptera (Sternorrhyncha and Auchenorrhyncha): an overview. Neotrop Entomol 30(4):501-516. https://doi.org/10.1590/S1519-566X2001000400001
Delabie JHC, Jahyny B (2007) A mirmecosfera animal: relações de dependência entre formis e outros animais. O Biológico, Sao Paulo 69:7-12.
Dias-Soares M, Correia IM, Santos JT,Delabie JHC, D’ávila S, Mariano CSF (2024) Facultative commensalism of gastropods (Mollusca: Gastropoda) in Neoponera verenae Forel, 1922 (Formicidae: Ponerinae) nests. Insect Soc https://doi.org/10.1007/s00040-024-00956-5
Diniz JLM, Brandão CRF, Yamamoto CI (1998) Biology of Blepharidatta ants, the sister group of the Attini: a possible origin of fungus-ants symbiosis. Naturwinssenschaften 85:270–274. http://dx.doi.org/10.1007/s001140050497
Eickwort GC (1990) Associations of mites with social insects. Annu Rev Entomol 35(1):469-488. https://doi.org/10.1146/annurev.en.35.010190.002345
Endo S, Itino T (2013) Myrmecophilous aphids produce cuticular hydrocarbons that resemble those of their tending ants. Popul Ecol 55:27-24. https://doi.org/10.1007/s10144-012-0355-0
Esteves FDA, Brandão CRF, Viegas K (2008) Subterranean ants (Hymenoptera, Formicidae) as prey of fossorial reptiles (Reptilia, Squamata: Amphisbaenidae) in centrl Brazi. Pap Avulsos Zool 48(28):329-334. https://doi.org/10.1590/S0031-10492008002800001
Eguchi K, Bui TV, Janssen R (2005) Gastropod guests (Prosobranchia: Pupinidae, and Pulmonata: Subulinidae) associated with the ponerine ant Diacamma sculpturatum complex (Insecta: Hymenoptera: Formicidae). Sociobiology 45:307–315
Fujihara RT, Forti C, Almeida MC, Baldin ELL (2011) Insetos de importancia económica: guía ilustrativo para identificação de Famílias. FEPAF, Botucatu
Glasier JRN, Poore AGB, Eldridge DJ (2018) Do mutualistic associations have broader host ranges than neutral or antagonistic associations? A test using myrmecophiles as model organisms. Insect Soc 65:639-648. https://doi.org/10.1007/s00040-018-0655-2
Goldsbrough CL, Shine R, Hochuli DF (2006) Factors affecting retreat-site selection by coppertail skinks (Ctenotus taeniolatus) from sandstone outcrops in eastern Australia. Austral Ecol 31:326-336. https://doi.org/10.1111/j.1442-9993.2006.01561.x
Hammer O, Harper D, Ryan P (2001) PAST: Paleontological statistic software for education and data analysis. Paleontologia Electronica 4:1-9
Hölldobler B, Kwapich CL (2019) Behavior and exocrine glands in the myrmecophilous beetle Dinarda dentata (Gravenhorst, 1806) (Coleoptera: Staphylinidae: Aleocharinae). PloS one 14(1):e0210524. https://doi.org/10.1371/jorunal.pone.0210524
Hölldobler B, Kwapich CL (2022) The guest of ants: how myrmecophiles interact with their hots. Belknap Press, Cambridge.
Hölldobler B, Wilson EO (1990) The Ants. Harvard University Press, Cambridge.
Hopkin SP, Read HJ (1992) The biology of Millipeds. Oxford University Press, Oxford.
Ivens ABF, von Beeren C, Blüthgen N, Kronauer DJC (2016) Studying the complex communities of ants and their symbionts using ecological network analysis. Ann Rev Entomol 61:353-371. https://doi.org/10.1146/annurev-ento-010715-023719
Janssen R, Witte V (2002) Allopeas myrmekophilos n. sp., the first snail reported as living in army ant colonies (Gastropoda: Pulmunota: Subulinidae). Archiv für Molluskenkunde 131:211-215.
Kistner DH (1982) The social insects bestiary. In: Hermann HR (ed) Social Insects, Academic Press, New York, pp 1-244
Kronauer DJ, Pierce NE (2011) Myrmecophiles. Current Biology 21(6):R208-R209.
Kruskal JB (1964) Multidimensional scaling by optimizing goodness of fit to a nonmetric hypothesis. Psychometrika 29:1-28. https://link.springer.com/article/10.1007/BF02289565
Lacau S, Fresneau D, Delabie J, Jahyny B, Montreuil O, Villemant C (2001) Uma nova associação entre as larvas mirmecófilas de suas espécies de Lampyridae (Insecta: coleoptera) e a formiga Typhlomyrmex rogenhoferi Mayr, 1862 (Formicidae, Ponerinae). Anasi do XV Econtro de Mirmcologia, IAPAR, Londrina, PR. 239-241.
Lapeva-Gjonova A (2013) Ant-associated beetle fauna in Bulgaria: a review and new data. Psyche 2013:242037. https://doi.org/10.1155/2013/242037
Lenhart PA, Dash ST, Mackay WP (2013) A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera: Formicidae) J Hymenopt Res 31: 119-164. https://doi.org/10.3897/JHR.31.4335
Lehtinen PT (1987) Association of uropodid, prodinychid, polyaspidid, antennophorid, sejid, microgynid, and zeconid mites with ants. Ent Tidskr 108:13-20
Lienhard C (1998) Psocoptères Euro-Méditerranéens. Faune de France 83. Fédération Francaise des Sociétés de Sciences Naturelles, Paris, France.
Lopes JMS, Oliveira AR, Delabie JHC (2015) Interações formigas/ácaros, com ênfase em ácaros foréticos associados a poneromorfas. In: Delabie JHC, Feitosa RM, Serrão JE, Mariano CSF, Majer JD (eds) As formigas Poneromorfas do Brasil, Editus,Ilhéus, pp 375-387
McGee KM, Eaton W (2014) The effects of the conversion of a primary to a secondary tropical lowland forest on bullet ant (Paraponera clavata) foraging behavior in Costa Rica: a possible indicator of ecosystem condition. J Insect Behav 27:206-2016. https://doi.org/10.1007/s10905-013-9413-5
McIver JD, Stonedahl G (1993) Myrmecomorphy: morphological and behavioral mimicry of ants. Annu Rev Entomol 38:351-377. https://doi.org/10.1146/annurev.en.38.010193002031
Mendonça CAF, Pesquero MA, Carvalho RDSD, de Arruda FV (2019) Myrmecophily and myrmecophagy of Attacobius lavape (Araneae: Corinnidae) on Solenopsis saevissima (Hymenoptera: Myrmicinae). Sociobiology 66(4):545-550. https://doi.org/10.13120/sociobiology.v66i4:4431
Migliorini GH, Ronque MU, Guipponi APL (2019) Stenochrus portoricensis (Arachnida: Schizomida) living in a nest of the fire ant Solenopsis saevissima (Hymenoptera: Formicidae) in the Atlantic forest, Brazil. Arachnology 18(2):127-128. https://doi.org/10.13156/arac.2018.18.2.127
Moleiro HR, da Silva-Melo A, Giannotti E (2021) Nest architecture and animals associated with Neoponera verenae (Forel) (Formicidae, Ponerinae). Sociobilogy 68(3):e6246. https://doi.org/10.13102/sociobiology.v68i3.6246
Moore W, Robertson JA (2014) Explosive adaptative radiation and extreme phenotypic diversity within ant nest beetles. Current Biology 24(20):2435-2439. https://doi.org/10.1016/j.cub.2014.09.022
Moreira I, Cruz CD, Fernandes AK, Delabie JHC, Castaño-Meneses G, Mariano C (2020) Estudo compartivo da fauna de comensais nos formigueiros de três espécies de grande tamanho da mirmecofauna brasileira (Hymenoptera: Formicidae). Bol Mus Para Emílio Goeldi 15(2):377-391. https://doi.org/10.46357/bcnaturais.v15i2.303
Mota-Filho TMM, Sousa KKA, Camargo RS, Oliveira JVLC, Caldanto N, Zeppelini D, Forti LC (2021) First record of Cyphoderus innominatus Mills, 1938 (Collembola: Paronellidae) in early colonies of the leaf-cutting ant Atta sexdens. Sociobiology 68(2):35922. https://doi.org/10.13102/sociobiology.v68i2.5922
Oliveira JVLC, Zeppleini D, Castaño-Meneses G, Palacios-Vargas JG (2023) Neotropical Cyphoderus (Collembola: Paronellidae), with comments about myrmecophily and the description of new species. Neotrop Entomol 52(4): 652-696. https://doi.org/10.1007/s13744-02201015-z
Oliveira MA, Della Lucia TMC (1993) Inquilinismo de Phylodryas olfersii (Reptilia, Squamta, Columbridae) em ninhos de Acromyrmex subterraneus (Hymenoptera, Formicidae, Attini). Rev Bras Entomol 37:113-115
Ostrovsky A, Georgiev D (2020) New Psocptera (Hexapoda, Insecta) records from Belarus. ZooNotes 157:1-3. https://doi.org/10.5281/zenodo.3753063
Palacios-Vargas JG, Mejía-Recamier BE, Zeppelini D (2013) Técnicas atuais para estudo de micro e mesoartrópodes de solo. Eduepb, Campina Grande
Parker J (2016) Myrmecophily in beetles (Coleoptera): evolutionary patterns and biological mechanisms. Myrmecol News 22:65-108. https://doi.org/10.25849/myrmecol.news_022_065
Parker J, Kronauer DJC (2021) How ants shape biodiversity. Curr Biol 31(19):R1208-R1214
Parmentier T (2020) Guests of social insects. In: Starr C (eds) Encyclopedia of social insects. ISBN: 978-3-319-90306-4 https://doi.org/10.1007/978-3-319-90306-4_164-1, Springer, Cham. Pp 1-15
Parmentier T, Claus R, De Laender F, Bonte D (2021) Moving apart together: co-movement of a symbiont community and their ant host, and its importance for community assembly. Mov Ecol 9: 25. https://doi.org/10.1186/s40462-021-00259-5
Peeters C, Hölldobler B, Moffet M, Musthak Ali TM (1994) “Wall-papering” and elaborate nest architecture in the ponerine ant Harpegnathos saltaror. Insectes Soc 41:211-218. https://doi.org/10.1007/BF01240479
Pérez-Lachaud G, Lachaud J-P (2014) Arboreal ant colonies as “hot-points” of cryptic diversity for myrmecophiles: the weaver ant Camponotus sp. aff. textor and its interaction network with its associates. PLoS One 9(6):e100155. https://doi.org/10.1371/journal.pone.0100155
Prado LP, Feitosa RM, Triana SP, Gutierrez JAM, Rousseau GX, Silva RA, Siqueira GM, Santos CLC, Silva FV, Silva TSR, Ferreira AC, Silva RR, Andrade-Silva J (2019) An overview of the ant fauna (Hymenoptera: Formicidae) of the state of Maranhão. Pap Avulsos Zool 59:3201995938. https://doi.org/10.11606/1807-0202/2019.59.38
Rees DP, Walker AJ (1990) The effect of temperature and relative humidity on population growth of three Liposcelis species (Psocoptera: Liposcelidae) infesting stored products in tropical countries. Bull Entomol Res 80(3):353-358. https://doi.org/10.1017/S0007485300050562
Rettenmeyer CW, Rettenmeyer ME, Joseph J, Berghoff SM (2011) The largest animal association centered on one species: the army ant Eciton burchellii and its more than 300 associates. Insectes Soc 58(3):281-292. https://doi.org/10.1007/s00040-010-0128-8
Rocha FH, Lachaud J-P, Pérez-Lachaud G (2020) Myrmecophiloud orgnisms associated with colonies of the ponerine ant Neoponera villosa (Hymenoptera: Formicidae) nesting in Aechmea bracteate bromeliads: a biodiversity hotspot. Myrmecol News 30:73-92. https://doi.org/10.25849/myrmecol_news_030
Rosa C (2008) Um estranho no ninho: efeito indireto da presença da aranha mirmecófaga Dipoena bryatae (Araneae: Theridiidae) no aumento da herbivoria em Hirtella myrmecophila (Chrysobalanceae). In: Machado G, Camargo JLC (eds) Livro do curso de campo Ecologia da Floresta Amazônica, PDBFF/INPA, Manaus,
Sánchez-Piñero F, Gómez JM (1995) Use of ant-nest debris by darkling beetles and other arthropod species in an arid system in south Europe. J Arid Environ 31(1):91-104. https://doi.org/10.1006/jare.1995.0052
Santos, P.P.; Vasconcellos, A.; Jahyny, B. & Delabie, J.H.C. (2010) Ant fauna (Hymenoptera; Formicidae) associated to arboreal nests of Nasutitermes spp. (Isoptera, Termitidae) in a cacao plantation in southern Bahia, Brazil. Revista Brasileira de Entomologia 54 (3): 450-454.
Schmidt JO, Overal WL (2021) Giant Amazonian ants (Dinoponera). In: Starr C (eds) Encyclopedia of social insects. ISBN: 978-3-319-90306-4 https://doi.org/10.1007/978-3-319-90306-4_164-1, Springer, Cham. Pp 434-439
Schönrogge K, Wardlaw JC, Peters AJ, Everett S, Thomas JA, Elmes GW (2004) Changes in chemical signature and host specificity from larval retrieval to full social integration in the myrmecophilous butterfly Maculinea rebeli. J Chem Ecol 30:91-107. https://doi.org/10.1023/B:JOEC.0000013184.18176.a9
Sierwald P (2007) MILLI-PEET: Illustrated key to Order. The Field Museum. https://www.fieldmuseum.org/science/special-projects/milli-peet-class-diplopoda/milli-peet-millipedes-made-easy/milli-peet-key. Accessed 26 June 2024
Silva VM, Moreira GF, Lopes JMS, Delabie JHC, Oliveira AR (2018) A new species of Cosmolaelaps Berlese (Acari:Laelapidae) living in the nest of the ant Neoponera inversa (Smith) (Hymenoptera: Formicidae) in Brazil. Syst Appl Acarol 23(1):13-24. https://doi.org/10.11158/saa.23.1.2
Sodano J, Oufiero CE, Schneider SA, LaPolla JS (2024) Scale insect (Hemiptera: Coccomorpha) morphology is transformed under trophobiosis. Ann Entomol Soc Am 117(1):49-63. https://doi.org/10.1093/aesa/saad033
Staniec B, Zagaja M (2008) Rove-beetles (Coleoptera, Staphylinidae) of ant nest of the vicinities of Lezajsk. Ann Univ Mariae Curie-Sklodowska Sect. Biol 63(1):111-127. https://doi.org/10.2478/v10067-008-0009-y
Sudd JH, Franks NR (1987) The behavioural ecology of ants. Springer Dordrecht. https://doi.org/10.1007/978-94-009-3123-7
Tierno de Figueroa JM, López-Rodríguez MJ (2019) Trophic ecology of Plecotera (Insecta): a review. Europ Zool J 86(1):79-102. https://doi.org/10.1080/24750263.2019.1592251
Triplehorn CA, Johnson NF (2011) Estudos dos Insetos.Cengage Learning, São Paulo
Tuma J, Eggleton P, Fayle TM (2020) Ant-termite interactions: an important but under-explored ecological linkage. Biol Rev 95(3):555-572. https://doi.org/10.1111/brv.12577
Vasconcellos A, Santana GG, Souza AK (2004) Nest spacing and architecture and warming of males of Dinoponera quadriceps (Hymenoptera: Formicidae) in a remnant of the Atlantic Forest in Northeast Brazil. Br J Biol 64:357-362. https://doi.org/10.1590/S1519-69842004000200022
Verdcourt B (2002) Two new species of Curvella Chaper (Gastropoda, Pulmonata, Subulinidade) from the East Usambara Mts., Tanzania. Basteria 66:107-112
Voigländer K (2011) 15 Chilopoda – Ecology. In: Minelli A (ed) Treatise on Zoology - Anatomy, Taxonomy, Biology. The Myriapoda, Vol. 1. Brill, Boston, pp 309-325 https://doi.org/10.1163/9789004188266_016
Whitfield SM, Donnelly MA (2006) Ontogenetic and seasonal variation in the diets of a Costa Rican leaf-litter herpetofauna. J Trop Ecol 22:409-417. https://doi.org/10.1017/S0266467406003245
Wilson EO (1971) The Insect Societies. Harvard University Press, London.
Wilson EO (2003) Pheidole in the New World. A dominant, hyperdiverse ant genus. Harvard University Press, London.
Witte V, Janssen R, Eppenstein A, Maschwitz U (2002) Allopeas myrmekophilos (Gastropoda, Pulmonata), the first myrmecophilous mollusc living in colonies of the ponerine army ant Leptogenys distinguenda (Formicidae, Ponerinae). Insect Soc 49:301-305. https://doi.org/10.1007/PL00012646

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

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