André Felipe de Araujo Lira

Spatio-temporal microhabitat use by two co-occurring species of scorpions in Atlantic rainforest in Brazil

With the increasing devastation of the tropical rain forest, there is a critical need to understand how animal forest communities are structured and how habitat degradation will affect these communities. We conducted a field survey to investigate the microhabitat preferences of two co-occurring species of scorpions (Tityus pusillus and Ananteris mauryi) in a fragment of Atlantic rainforest, as well as their abundance and their ecological niche, during both the dry and rainy seasons. Behavioural aspects related to the use of the environment and the proportions of juveniles and adults are also described. The occurrence of intra- and interspecific coexistence was assessed by active search. In addition, pitfall catches were used to assess the structure of the population in the dry and rainy seasons. The differential patterns of spatial distribution in the litter layers provided evidence of partial niche partitioning between the two coexisting scorpion species depending on age and climatic conditions. Abundance, foraging behaviour and age structure (juveniles and adults) were seasonally influenced. We conclude that the diverse and subtle behaviours involved in interaction and habitat use may facilitate species coexistence. Resource partitioning and refuge sharing on a temporal and/or spatial scale, as well as predation pressure, may drive the dynamics and spatial distribution of scorpion species in the rain forest environment.

Diversity of scorpions (Chelicerata: Arachnida) in the Atlantic Forest in Pernambuco, northeastern Brazil

This study was conducted to measure the biodiversity of scorpion species in the remnants of the Brazilian northeastern Atlantic Forest, an important center of biodiversity and endemism. Collections were performed in twelve forest fragments in Sirinhaem municipality, Pernambuco State, through active searches at night by using UV lamps between 19:00h and 21:00h during the new moon phase in December 2012 and January 2013. A total of 1,125 specimens from two genera and five species were collected: Tityus pusillus Pocock, 1893; Ananteris mauryi Lourenco, 1982; Tityus brazilae Eickstedt & Lourenco, 1984; Tityus neglectus Mello-Leitao, 1932; and Tityus stigmurus (Thorell, 1876), all belonging to the family Buthidae C.L. Koch, 1837. The most abundant species was T. pusillus (90.7%), followed by A. mauryi (7.1%). Tityus brazilae , T. neglectus, and T. stigmurus together represented less than 3% of the individuals sampled.

The effect of habitat fragmentation on the scorpion assemblage of a Brazilian Atlantic Forest

Habitat fragmentation is a topic widely studied in ecology; however, its effects on the assemblage of the order Scorpiones is less well understood. Aiming to fill this gap, this study assessed the effect of fragmentation on the assemblage of these arachnids in 12 Brazilian Atlantic forest fragments. Five environmental variables were measured (depth and dry mass of litter, understory density, canopy openness, and diameter at breast height of the trees), and the fragment area, vegetation cover, connectivity and elevation assessed. The animals were collected during the dry season and, identified at night with the use of ultraviolet light lamps. The analyzed scorpion assemblage in the landscape was characterized by the species Tityus pusillus, T. stigmurus, T. neglectus, T. brazilae, and Ananteris mauryi, with a maximum of three species co-occurring per fragment. Only the fragment size and the dry mass of litter showed a positive relationship with the composition of scorpions. These results suggest that the habitat of scorpions responds to environmental attributes and landscape metrics at both higher (fragment size) and lower (leaf litter) scales. Our study was able to expand our knowledge of how scorpions respond to habitat changes in the Atlantic Forest. We conclude that fragmentation and habitat quality are determining factors for the assemblage of these arachnids.

Dominant Dung Beetle (Coleoptera: Scarabaeidae: Scarabaeinae) Species Exhibit Wider Trophic Niches on Fruits, Excrement, and Carrion in Atlantic Forest, Brazil

Dung beetles of the subfamily Scarabaeinae encompass an important part of the copronecrophagous fauna in the tropical and temperate regions (Hanski and Cambefort 1991). These insects exert extremely important environmental services, acting on recycling soil nutrients, outcompeting and controlling cattle parasites, and serving as efficient bioindicators (Nichols et al. 2008). Neotropical forests contain the richest copro-necrophagous dung beetle fauna of the world, and they are where frugivory in this group is most studied (Halffter and Halffter 2009). Elevated competition occurs among Neotropical dung beetles, associated with the high diversity of dung beetles and relative scarce availability of large mammal droppings in this region (Hanski and Cambefort 1991; Halffter and Halffter 2009). This pattern is a result of historically low dung availability, leading to different strategies of resource partitioning (Halffter 1991; Halffter and Halffter 2009). Species can develop in order to exhibit singular adaptations of feeding patterns, presenting generalist habits as a response to dung scarcity (Martín-Piera and Lobo 1996). Utilization of a variety of resources reduces the competition among species, permitting the occurrence of higher diversity in environments (Hanski and Cambefort 1991; Halffter and Halffter 2009). The aim of the present study was to investigate the attractiveness of dung beetles for excrement, carrion, and rotten fruits in a fragment of Atlantic Forest. We hypothesize that species that use a wider diversity of resources will be more abundant, since dung beetles with a more generalist diet tend to exhibit higher success in feeding, sexual maturation, and consequential reproduction (Filgueiras et al. 2009; Silva et al. 2012). The study was conducted in December 2012 in a fragment of northeastern Atlantic Forest (8°33′50.2′′ S 35°10′03.4′′ W) in the municipality of Sirinhaém, Pernambuco, Brazil. The fragment is composed of ombrophilous, dense forest vegetation and has an area of 288 ha inserted in a matrix of sugarcane monoculture. To capture the dung beetles, 50 pitfall traps were installed with one of five bait types (50 g of bait per trap): carrion (bovine rotten spleen); human excrement; rotten fruits of cashew (Anacardium occidentale L.; Anacardiaceae); rotten fruits of banana (Musa sp.; Musaceae); and rotten fruits of mango (Mangifera indica L.; Anacardiaceae). Pitfall traps consisted of a plastic container (9.5 × 20 cm) buried in soil at the surface level, with a circular disk above it to protect it from rainwater and litter entrance. The baits were inserted into a small container attached to the plastic container. To maintain the dung beetles inside the pitfall, a solution of 200 ml of water with 10 g of salt and 10 ml of detergent were utilized inside the pitfall traps. Sets of 10 traps containing one bait type in each were placed in the field, totaling five groups and 50 traps. Traps with the same bait type were installed in a linear transect, each trap 10 m distant from another. The transects were installed in a straight line, each transect line separated by 50 m from the nearest

Distinct edge effects and reproductive periods of sympatric litter-dwelling scorpions (Arachnida: Scorpiones) in a Brazilian Atlantic forest

Edge effects have drastically affected species living in tropical forests. However, understanding how species respond to edge effects remains a challenge, owing to the many factors involved and different responses of each species thereto. Here, we analyzed how the abundance of two sympatric scorpion species (Tityus pusillus and Ananteris mauryi) and their potential prey varied as a function of microhabitat changes (litter depth, dry mass, and leaf shape) from edge to interior forest habitats. We further analyzed the contribution of potential prey to scorpion abundance and reproductive periods. Data were collected monthly at three 300-m² transects/site at distances of 10, 100, and 200m from the forest edge in a fragment of the Atlantic forest in northeastern Brazil, between April 2016 and March 2017. Scorpions responded differentially to edge effects, with A. mauryi abundance being similar along the edge-interior gradient, whereas T. pusillus had a higher abundance in the interior. As T. pusillus inhabit the top layer of the leaf litter, this species will possibly be more influenced by edge effects. In contrast, being a humicolous scorpion and inhabiting the bottom layers of leaf litter, A. mauryi would not be influenced by edge effects as much as T. pusillus. The reproductive period also was distinct between the two species, with T. pusillus reproducing in the dry season and A. mauryi in the rainy season. The oscillation in the abundance of different groups of prey at different periods maintained the overall prey abundance at a relatively constant level throughout the year, mitigating the effects of prey availability on the abundance and reproductive period of the scorpions. These results suggest that microhabitat exploitation is a key factor to sustain litter-dwelling scorpions in disturbed forest remnants and that T. pusillus can be an ecological indicator of edge effects.

Linking scorpion (Arachnida: Scorpiones) assemblage with fragment restoration in the Brazilian Atlantic Forest

ABSTRACT Habitat loss due to forest degradation can induce changes in species richness due to variation in species susceptibility to environmental stress. This is particularly important for species with highly specific microhabitats, such as scorpions that inhabit forest habitats. In this study, the richness and abundance of these arachnids were compared between an old-growth (mature) and secondary (65 years under natural restoration) forests. Seasonal influence was also evaluated by comparing diversity between dry and wet seasons. The animals were collected through nocturnal active search using UV lamps and pitfall traps in both areas (old-growth and secondary). Both environments showed similar breast heights of trees, litter depth, litter dry mass, and understory density, indicating a high level of restoration. Scorpion diversity (characterized by Tityus pusillus, T. neglectus, T. brazilae, Bothriurus asper, and Ananteris mauryi) and abundance were not influenced by the different historical usage of both areas. In contrast, the abundance of these arachnids was highly affected by rain regimes, and increased during the dry season. These results suggest that 65 years was a sufficient time period for restoration, making it possible to maintain similar scorpion assemblages in both environments.

Leucism in Tityus pusillus (Scorpiones: Buthidae): Report of a rare event in scorpions

Abstract Leucism is a congenital disorder in which the individual is born with partial hypopigmentation. It is quite common in vertebrates, but rare in invertebrates, especially in arachnids like scorpions. This paper presents the first record of this congenital disorder to be observed in the order Scorpiones. During field studies in the Area de Conservação Aldeia-Beberibe, a set of Atlantic forest fragments of 31,634 hectares, we collected a pregnant leucistic female Tityus pusillus Pocock, 1893. In this female, the variegated pattern described for the species was a lighter color than normal. The animal produced 10 normal juveniles (not leucistics). In addition, we analyzed 1,164 specimens from 17 populations deposited in the CA-UFPE to verify the frequency of leucism; there were no scorpions with leucism within the analyzed populations. Thus, a break in variegated pattern, as with the leucism described in this study, may increase the mortality rate due to predation.

Records of Perching by Dung Beetles (Coleoptera: Scarabaeidae: Scarabaeinae) in the Northeastern Atlantic Forest of Brazil

Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) are an important group of copronecrophagous beetles in the tropical and temperate regions (Hanski and Cambefort 1991). They are essential to the ecosystem because they cycle nutrients, disperse secondary seeds, and control populations of cattle parasites (Nichols et al. 2008). The Neotropical region is one of the most relevant regarding dung beetle diversity. However, it has a relatively low availability of mammal dung (one of the most important resources for these beetles) compared, for example, to the Ethiopian region (Hanski and Cambefort 1991; Halffter and Halffter 2009). In part, this may be a driver of high interspecific competition for food resources among dung beetles in the Neotropics (Gill 1991; Halffter et al. 1992; Davis et al. 2001). The high degree of competition for food resources may also lead to the development of adaptive strategies that reduce competition. The occupation of specific microhabitats, different patterns of feeding adaptations, and segregation of daily activities are some examples of strategies that dung beetle species employ to avoid high competition for resources (Halffter and Mathews 1966; Hernández 2007; Lopes et al. 2011; Salomão et al. 2014; Noriega 2015). Some dung beetle species perch (Howden and Nealis 1978; Louzada 1998). Perching is a less known but apparently highly important activity. This behavior has been observed in various biomes such as tropical, subtropical, temperate, and scarp forests (Davis 1999; Ballerio 2008; Jacobs et al. 2008; Medina and Campos 2013). Nevertheless, to the best of our knowledge, there are no records of such behavior for species in Pernambuco’s endemism center situated in the northeastern Brazilian Atlantic Forest. This ecosystem is considered one of the most important ecosystems in the world and extremely important for worldwide conservation efforts (Myers et al. 2000; Tabarelli et al. 2010). In the Atlantic Forest, dung beetles display a wide variety of strategies to reduce the competition for resources. They have specialized feeding and habitat occupation strategies (Forti et al. 2012; Costa et al. 2013; Salomão et al. 2014). Canthon (Canthon) virens Mannerheim, 1829, for example, preys on queen ants of the genus Atta Fabricius, 1805 during the ants’ nuptial flights (Forti et al. 2012). Another example of adaptation is the specialization on sloth dung by some species of the genus Trichillum Harold, 1868 and Uroxys Westwood, 1842 (Ratcliffe 1980). Based on such specializations, we can better understand some dung beetles’ adaptations to certain environments as they show strong niche segregations. In the present study, we present the first record of perching observed in dung beetles in fragments of the northeastern Brazilian Atlantic Forest. The observations of perching were made in situ during expeditions conducted in the morning (08:00-12:00 hr) and one expedition in the afternoon (15:00-16:00 hr) in four Atlantic Forest remnants (Table 1) in the State Pernambuco, northeastern Brazil. Perching beetles were collected and perching height above ground was measured using a metric tape. Body lengths of the specimens of Canthon Hoffmannsegg were DOI.org/10.1649/0010-065X-70.4.903

Metasomal Autotomy in Ananteris mauryi Lourenço, 1982 (Scorpiones: Buthidae)

Most animals possess defense mechanisms, which they use against their potential predators, including morphological and behavioral adaptations that reduce their chances of encountering a predator or increase the likelihood of surviving an attack (Edmunds 1974). Among these mechanisms, the autotomy of appendages is widely known in many taxa of both invertebrates and vertebrates (Maginnis 2006; Fleming et al. 2007). Autotomy is an efficient means of escaping from predators that grasp their prey (Formanowicz 1990; Arnold 1994; Klawinski and Formanowicz 1994), and increases the survival rates of the prey. However, the loss of an appendage in invertebrates, includes a wide variety costs, from almost no costs in some harvestmen (Guffey 1998), to locomotory costs and susceptibility to a subsequent predatory attack, decreasing the reproductive success, which in turn reduces their intraspecific competitiveness (Uetz et al. 1996; Fleming et al. 2007). J Insect Behav (2014) 27:279–282 DOI 10.1007/s10905-013-9431-3

Phoresy by Americhernes Aff. Incertus (Pseudoscorpiones: Chernetidae) on a Tropical Fly Fannia canicularis (Diptera: Fanniidae) in a Fragment of the Atlantic Forest, Brazil

ABSTRACT Pseudoscorpions employ phoresy for dispersal, in which one animal attaches to another for transportation. This study documents the occurrence and phoresy by Americhernes aff. incertus (Pseudoscorpiones: Chernetidae) on the fly Fannia canicularis (Diptera: Fanniidae) in a fragment of semi-deciduous seasonal Atlantic Forest, in the state of Pernambuco, northeast Brazil. The pseudoscorpions were found attached to the ventral surface of six individual flies. To our knowledge, no previous studies have shown the presence of Americhernes aff. incertus in the northeast Atlantic Forest and its use of flies as a dispersal method.