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Because of the exoskeleton, arthropods must have flexible areas to be able to move. Such regions are called arthrodial membranes and are particularly vulnerable to bacteria and fungi. Here, we analyzed the secretion in the glands underneath it in a Neotropical harvester (Arachnida, Opiliones) and tested whether it has antiseptical properties. We punctured the membrane, collected and diluted the secretion and quantified proteins and peptides in a spectrophotometer. We also fractionated and analyzed the samples in reversed-phase high-performance liquid chromatography (RP-HPLC) and then incubated the treated fractions and determined growth inhibition by measuring absorbance. The secretions resulted in 42 fractions, among which two had activity against the Gram-positive bacteria Micrococcus luteus and against the yeast Candida albicans. The low concentrations at which the secretions were active are relevant from a biotechnological point of view. For the animals, the secretions possibly prevent infections, including when they are attacked in these regions by predators that pick that spot to bite.

The subfamily Gonyleptinae is the second largest in Gonyleptidae, harboring over 100 species. Gonyleptinae is polyphyletic, nestled in the clade K92, and despite its richness, several species of that subfamily have not had their chemicals of the defensive secretions analyzed. Among these are Gonyleptes curticornis (Mello-Leitão, 1940) and G. horridus Kirby, 1819, the latter being particularly important because it is the type species of the genus, which in turn names the subfamily. Gonyleptes horridus is also used in many phylogenetic analyses, be it using morphological or molecular data. The chemical study of the secretions of these two species by GC-MS and NMR 1H showed the presence of 1-(6-isopropyl-3,4-dihydro-2H-pyran-2-yl)-methylbutanone, 1-(6-isopropyl-3,4-dihydro-2H-pyran-2-yl) isobutanone and 4- methyl-1-hepten-3-one in both species. On the other hand, 4-methyl-1 hexen-3-one, benzaldehyde and 3-octanone were observed only in G. curticornis. Both species are Gonyleptinae and chemical mapping of the group corroborates that vinyl ketones are synapomorphy of K92 and that the compound 1-(6-isopropyl-3,4-dihydro-2H-pyran-2-yl) isobutanone is synapomorphic to G. curticornis and G. horridus, but homoplastic to the genus Sodreana Mello-Leitão, 1922. 1-(6-(1-methyl-propyl)3,4-dihydro-2H-pyran-2yl)2-methylbutanone and 4-methyl-1-hepten3-one is also synapomorphic to G. curticornis and G. horridus, but homoplastic in Moreiranula saprophila.

Assessing information about prey before attacking is crucial for predators. Whip spiders (Arachnida, Amblypygi) are predatory arachnids that prey mainly on arthropods and whose first pair of legs concentrates many sensory structures. Nonetheless, these aspects of amblypygid biology have been mostly studied in a few species of Phrynidae and basic information about predatory behavior and sensory morphology remain scarce for other families. In this study, we described the sensory morphology of the antenniform leg and the predatory behavior of Charinus asturius Pinto-da-Rocha, Machado and Weygoldt, 2002, a representative of the family Charinidae. We report the presence of most sensilla types described in the better studied Phrynidae and highlight novel aspects such as the morphology of the tarsal organ and the reduced number of rod sensilla. Our behavioral results underscore the importance of antenniform legs in the predatory behavior of C. asturius. Finally, we suggest new avenues to advance the knowledge about the importance of antenniform legs for capturing living prey.

Consistent inter-individual differences in behavior have been shown in several animal groups, ranging from vertebrates to invertebrates. One of the most studied personality traits in animals is boldness, which is the tendency to expose to risky situations. Theory proposes that individuals’ state (e.g., body energy) would influence the expression of personality traits. In this study, we tested if boldness levels of the harvestman Mischonyx cuspidatus (measured as duration of death feigning/freezing after simulated predator attack) differ between two different states, namely sated and food deprived. We also tested if the degree of repeatability in boldness is affected by the individual state. We found no differences in the levels of boldness expressed by M. cuspidatus when comparing between different conditions (sated and food deprived) at a population level. However, we found that individuals showed more consistency in boldness when sated relative to a food-deprived condition. Finally, we suggest new avenues for future studies addressing personality in harvestmen.

Scorpionism is a worldwide problem that has already made thousands of victims, and multi-disciplinary approaches for controlling their populations are to be more successful. Hens are often mentioned as tools for controlling scorpions; however, systematic/experimental behavioral studies are not available. Moreover, there is no systematic information on the effect of scorpion venoms on hens. Using the venomous yellow scorpionWe attracted hens with corn powder, offered them scorpions and then recorded the hen-scorpion interaction. To test the effects of the sting we manually removed the scorpion's telson.We found that some hens ate up to six scorpions within minutes. By means of an ethogram and drawings, we showed that they exhibited several aversive behaviors when capturing scorpions. Removal of the scorpion telson stopped the aversive reactions, which was not observed in the control group. Finally, hens did not exhibit atypical behaviors after 1, 7 and 30 days and were all alive after 30 days.This is the first empirical and video recorded study providing evidence that hens are clearly affected by scorpion venom but do not die. Therefore, they may have potential to be used in biological control of these arthropods.

Animals that live by rivers may benefit from being able to cross them, but behavioral adaptations are needed. Additionally, being able to remain submerged is also important if the animal moves under water. Here we asked whether the harvestman Heteromitobates discolor (Opiliones), that lives by rivers, (a) can propel itself across the water surface, (b) moves into the water if disturbed and (c) can survive for long periods when submerged. Heteromitobates discolor exhibited two gaits on water, whereas a strictly terrestrial species was not able to propel itself. When experimentally submitted to simulated predator attack on a rock on the river, H. discolor walked onto the water, while a strictly terrestrial species did not. Finally, it was able to survive for 6 h under water, presumably due to the conspicuous air film that formed around its body, which was also observed in a strictly terrestrial species. Altogether, these observations suggest that the aquatic environment is not a barrier for regular activity and can be used as an extension of the terrestrial environment for H. discolor.

Sexual differences in morphology can evolve by sexual selection and/or natural selection. In some species, only males have morphological structures that are used as weapons. Since some weapons may also be used for defensive purposes, males and females may behave differently towards predators. In some species of harvestmen (Arachnida and Opiliones), males have sharp apophyses (“spines”) on their 4th pair of legs whereas females lack them. Those apophyses are used in male–male fights and in antipredatory behaviors. The harvestmen antipredatory repertory also encompasses passive defenses such as thanatosis (death feigning), retaliation (attack on predators), and chemical defense. Due to the sexual differences on weaponry, we hypothesized that males and females of Mischonyx cuspidatus (Gonyleptidae) rely on different defensive strategies. We experimentally induced males and females to perform 3 defensive behaviors: thanatosis, pinching with legs, and chemical release. We predicted that females would engage more in passive and chemical defenses than males, whereas males would rely more on retaliation than females. As expected, females performed thanatosis more often than males. Likewise, males performed retaliation more often than females. We did not find differences in the rate of chemical defense use between the sexes. This study provides evidence that due to sexual dimorphism, alternative antipredatory behaviors may have been selected in the different sexes in M. cuspidatus.

Animals use external and/or internal cues to navigate and can show flexibility in cue use if one type of cue is unavailable. We studied the homing ability of the harvestmanHeteromitobates discolor(Arachnida, Opiliones) by moving egg-guarding females from their clutches. We tested the importance of vision, proprioception, and olfaction. We predicted that homing would be negatively affected in the absence of these cues, with success being measured by the return of females to their clutches. We restricted proprioception by not allowing females to walk, removed vision by painting the eyes, and removed the odours by removing the clutch and cleaning its surroundings. We found that vision is important for homing, and in the absence of visual cues, proprioception is important. Finally, we found increased homing when eggs were present, and that the time of the day also influenced homing. We highlight vision as a previously overlooked sensory modality in Opiliones.

Most harvestman species are dependent on high humidity levels and amenable temperatures for homeostasis. While they are known to actively choose environments with these conditions, no thermo-/hygroreceptor has yet been identified in harvestmen. Using electron microscopy, we investigated the ultrastructure of two types of hair sensilla of the armored harvestman Heteromitobates discolor (Laniatores, Gonyleptidae): namely the sensillum basiconicum and the hooded sensillum. Both structures occur in small numbers (sensilla basiconica: 28 units; hooded sensilla: 4 units) and are distributed on the distal parts of the legs. On the distalmost tarsomeres I and II, the receptor cells of paired sensilla basiconica and single hooded sensillum occupy a large volume of the inner tissue, and appear tightly associated. The sensillum basiconicum is innervated by 3-4 dendrites and has a longitudinal slit giving the impression of a shaft with two flaps, resembling a beak. The slit probably allows for evaporation of sensillum lymph. The hooded sensillum is innervated by two bundles of three dendrites each, has two pore-like structures on its tip and displays an unusual reticulate cuticle of the shaft. Details of shaft cuticle, the evidence of evaporation of sensillum lymph, and specific innervation patterns support our hypothesis that sensilla basiconica are thermo- and/or hygroreceptors. Even though the definite function of hooded sensilla remains unclear, its putative receptor modalities are assessed by means of specific ultrastructures. Finally, we discuss with regard to functional ultrastructures as to whether the evaporation system of sensilla basiconica fits mechanisms of hygroreception as known from current literature.

Pedipalps in laniatorid harvestmen are usually cylindrical and raptorial, but species in Cosmetidae are exceptional in that adults, but not immature, have pedipalps flattened as a spoon. Th...

Phylogenetic relationships in Opiliones (Arachnida) at the suborder level have greatly stabilized in recent years, largely due to advances in molecular systematics. Nonetheless, identifying morphological characters in the context of well-resolved phylogenies is essential for testing new systematic hypotheses and establishing diagnostic markers. Here, we investigate with SEM a promising character system across Opiliones: the sensilla on the distalmost article of legs I and II. We identified four discrete characters and scored species of nearly all families of Laniatores (28 families, 44 species), three Dyspnoi, two Eupnoi and two Cyphophthalmi. Using a phylogenetic backbone compiled from recent and ongoing phylogenomic studies, we trace the evolution of these sensilla using ancestral state reconstruction. We discover a widespread occurrence of three sensilla (a pair of sensilla basiconica and one hooded sensillum) on the anterior legs of all families of Laniatores studied, and that comparable structures occur in the other suborders of Opiliones. Our analysis shows that this sensory field provides diagnostic information at different levels of phylogenetic relationships. We discuss the implications of the widespread occurrence of these sensilla in Opiliones, which have recently been hypothesized as hygro-/thermoreceptors and their putative homology with tarsal organs in Arachnida.

Prey morphology and size are known to influence a predator’s decision to attack and consume particular prey; however, studies that evaluate both traits simultaneously are uncommon. Here, we first described the trophic niche in the mygalomorph spider Paratropis sp. These spiders have a narrow trophic niche and feed mainly on sympatric species such as larvae of lepidopterans and of beetles such as carabids, passalids and scarabeids. Second, we evaluated the effect of prey taxon and size on acceptance and immobilization duration, and built an ethogram of the predator’s behaviors. For each prey taxa, we offered large (approximately same size than the spider) and small larvae (approximately half of the size of the spider) of the aforementioned prey. We classified carabid beetle larvae as the most dangerous prey because of their sharp mandibles and predatory habits, followed by scarabeid larvae; lepidopteran and passalids larvae were considered to be non-dangerous prey. We did not find a significant effect of prey taxon or size on spiders’ acceptance. Prey size did not affect the time invested on each behavioral category, but prey taxon did. Moreover, although spiders used a similar strategy for capturing prey, they spent more time biting carabid larvae than other prey. Our results suggest that, at least in our study area, prey that are dangerous or the predator’s size do not seem to affect Paratropis sp. acceptance per se, but can cause the predator to change the time budget allocated to each behavior.

Acquiring food requires success in all the distinct phases of foraging, among which are detecting, capturing and handling prey. We have looked at prey detection, capturing and handling in three species of armored harvestmen differing in leg length and pedipalp morphology: Discocyrtus pectinifemur, Heteromitobates discolor and Gryne perlata. We recorded males and females in captivity capturing 0.5- to 0.7-mm-long immature crickets without legs III and provide the first detailed description of prey capture in harvestmen of the suborder Laniatores. We have shown that these three species can detect live prey without touching it but only at close range (

There are records of glands that produce sexual pheromones that are released into the environment or applied directly on sexual partners. Within Opiliones (Arachnida), several harvestmen in the suborder Laniatores have sexually dimorphic glands on legs I and IV, the mode of use of which is recorded only in two species but their function is unknown: while walking, males rub the glands against the substrate or against their body. Here we test an alternative and non-exclusive hypothesis that the glands present on the legs of male Gryne perlata (Cosmetidae) produce contact pheromones used in mating. We predicted that males would touch the females with the gland openings or with other male body parts previously rubbed by these glands. We also predicted that there are chemoreceptors on those parts of the females where males touch them. We analyzed 13 videos of G. perlata mating, a species in which the males have glands on legs I and IV of unknown function. We also analyzed 14 videos of Discocyrtus pectinifemur (Gonyleptidae) mating as a control, a species that lacks these glands. Finally, we looked for chemoreceptors on the legs of female G. perlata using a scanning electron microscope. During copulation, males of both species rubbed the legs of females with their first pair of legs, but not with the regions of these legs where the openings of the glands are. The fourth pair of legs were only used to support the body. Rubbing other body parts of the female by males with their glands was not observed during mating. Setae on the legs of the female did not have tip pores and therefore do not seem to be chemoreceptors. We therefore did not find any evidence that these sexually dimorphic glands in G. perlata release contact pheromones during mating.

Chemoreception is a main sensory modality in harvestmen but there has been no experimental evidence of attraction to conspecifics by olfaction excluding contact chemoreception as an alternative explanation. Here, using a Y-maze olfactometer we tested whether the harvestman Mischonyx cuspidatus (Roewer, 1913) is attracted to volatiles of conspecifics. We found that males were attracted to volatiles of other males, but not females. Females were not attracted to female's volatiles, but were marginally more often attracted to volatiles of males. Our results help understanding mechanisms in mating and social behavior in the order Opiliones.

Sexually dimorphic traits are widespread in animals, and include sex-specific weapons, ornamentation and, although less noticed, glands and associated structures. In arachnids, certain lineages of the order Opiliones exhibit diverse forms of dimorphism in the armature and length of appendages (common in Laniatores), as well as in the presence of sexually dimorphic glands (mostly investigated in Cyphophthalmi), positing harvestmen as promising models to study sexual dimorphism. Whereas the evolution and ecological significance of armature have been the focus of recent attention, sexually dimorphic glands remain understudied in groups other than Cyphophthalmi, despite being widespread in Opiliones. We therefore selected the amphi-Pacific family Zalmoxidae as an ideal taxon to investigate the evolutionary dynamics of this trait. We first describe four new species of Palaeotropical Zalmoxis, including a species with sexually dimorphic glands, and describe the morphology of zalmoxid species with sexually dimorphic glands using scanning electron microscopy. Using a previously assembled six-locus dataset supplemented with new terminals, and applying stochastic character mapping, we infer that sexually dimorphic glands evolved once in the Neotropics and at least four times in the Palaeotropic zalmoxids, revealing the evolutionary lability of this trait.

Sexually dimorphic traits are widespread in animals, and include sex-specific weapons, ornamentation and, although less noticed, glands and associated structures. In arachnids, certain lineages of the order Opiliones exhibit diverse forms of dimorphism in the armature and length of appendages (common in Laniatores), as well as in the presence of sexually dimorphic glands (mostly investigated in Cyphophthalmi), positing harvestmen as promising models to study sexual dimorphism. Whereas the evolution and ecological significance of armature have been the focus of recent attention, sexually dimorphic glands remain understudied in groups other than Cyphophthalmi, despite being widespread in Opiliones. We therefore selected the amphi-Pacific family Zalmoxidae as an ideal taxon to investigate the evolutionary dynamics of this trait. We first describe four new species of Palaeotropical Zalmoxis, including a species with sexually dimorphic glands, and describe the morphology of zalmoxid species with sexually dimorphic glands using scanning electron microscopy. Using a previously assembled six-locus dataset supplemented with new terminals, and applying stochastic character mapping, we infer that sexually dimorphic glands evolved once in the Neotropics and at least four times in the Palaeotropic zalmoxids, revealing the evolutionary lability of this trait.

The study of sensory structures has the potential to provide insights into the natural history and evolution of animals. The sensory structures of arachnids are usually concentrated on the pedipalps (the tritocerebral appendages) or on the distal podomere (tarsus) of the anterior walking legs, the latter being the case for armored harvestmen (Opiliones, Laniatores). Therefore, modifications of the tarsus could have direct impacts on the sensory equipment of these animals. Using scanning electron microscopy, we investigated the sensory equipment in an extreme case of reduction in tarsal articles in the harvestman Sandokan truncatus (Sandokanidae), which bears a single tarsomere in all legs, and the potential consequences of this reduction. Additionally, we review the literature on the natural history of the family Sandokanidae. Tarsomeres of all legs are equipped with gustatory sensilla, mechanoreceptors, and a pore organ, but wall-pored olfactory sensilla are restricted to tarsi I and II. Tarsi II present a higher density of olfactory sensilla and also putative campaniform sensilla (strain detectors), which indicates a special sensory function of this pair of legs. Other podomeres are covered with shelled sensilla, a probable chemoreceptor previously unreported in Opiliones. Overall, S. truncatus has types of sensilla largely comparable to harvestmen with longer and subdivided tarsi. However, S. truncatus also exhibits extra-tarsal sensory fields of sensilla basiconica (putative thermo-/hygroreceptors) in previously undescribed sites, and the unique pore organs. Our results establish a basis for further research investigating the natural history, as well as the evolutionary correlations and mechanistic causes of the tarsal reduction in this enigmatic lineage.

We provide a first scanning electron microscopy examination of the Paleotropical harvestman family Podoctidae (Opiliones: Laniatores), focusing on the distitarsus of the legs of Metibalonius sp. Distitarsi I and II are mostly equipped with olfactory sensilla chaetica with wall pores, while those of legs III and IV have gustatory sensilla chaetica with a tip pore, ventral trichomes with ovate tips (non-sensory) and a type of spatulate seta. Spatulate setae are present in adults of both sexes, with no apparent sexual dimorphism, but they are absent in the nymph. Seven of these setae are inserted on the frontal surface of the last tarsomere of legs III and IV, with the tips oriented ventrally. Each seta has an s-shaped socketed shaft, which terminates distally in a spatula-shaped structure. The distribution of spatulate setae, restricted to legs III and IV (walking legs), the position on the distitarsi, and the typical spatulate shape suggest an adhesive function for these structures. Morphology and position suggest that the socketed spatulate setae of Metibalonius sp. and the previously reported scopular spatulate setae of other harvestmen constitute two distinct types of adhesive structures, highlighting the diversity of adhesive structures in Laniatores. Future investigations about the natural history of this species and internal morphology of spatulate setae are necessary to test further functional hypotheses and to determine their behavioral role.

Preliminary observations suggested that the Neotropical harvestman Jussara sp. (Sclerosomatidae, Eupnoi) had a patchy distribution and a marked seasonality, comparable to species in the Northern hemisphere. This would be an unusual pattern, since conditions of temperature and humidity are very different between tropical and temperate environments. Using a mark-recapture method, we investigated the phenology, habitat use and individual movement of Jussara sp. in a tropical rainforest in southern Brazil. We found a very marked seasonality in the population of adults, with a positive correlation between temperature, humidity and number of adults found. Adults were found only in the wet and warm months and significantly preferred the plant Psycothria suterella (Rubiaceae) as a resting substrate, spending the days motionless on its leaves, often in groups. The same individuals were found for a maximum of three months after their first capture, 30 m from their original site of capture. At night, the ha...

Prey consumption depends on the predator's ability to locate, capture and handle prey. We investigated these three steps in interactions between the delicate-bodied recluse spider Loxosceles gaucho (Araneae) and a heavy-bodied and armoured harvestman, Mischonyx cuspidatus (Opiliones). Although previous research suggested that the hard integument of such harvestmen protects them from being preyed upon by spiders larger than Loxosceles , indirect evidence suggested that Loxosceles spiders can subdue these prey. In the present study, we tested the following three hypotheses with regard to L. gaucho : (1) spiders use chemical cues left by prey to select foraging sites; (2) vibratory cues of prey are essential information in the predatory process; and (3) the spider's web sheet allows adequate handling of prey so vulnerable body regions of the prey can be bitten. To understand how a delicate predator can overcome the defences of a heavy-bodied and well-defended prey, we also quantitatively described the spider's behaviour. To test hypothesis 1, we compared the time spent in areas with harvestmen, crickets and no cues. For hypothesis 2, we compared latency to bite and number of bites in the presence or absence of vibratory information, and for hypothesis 3, we compared latency to detect prey, latency to capture prey and predation success. All three hypotheses were rejected. Loxosceles gaucho seems to be exceptional among spiders by not needing its web, indirect prey chemical cues, or prey's substrate borne vibrations to hunt the tested prey. What enables L. gaucho to prey upon M. cuspidatus is its unique hunting strategy, compared to previously studied spiders: it touches the prey with its tarsi possibly to locate weak parts of the prey's body, such as joints and distal parts of the legs, then it delivers several bites to these vulnerable areas. Our study is the first to document and describe how recluse spiders overcome the defences of an armoured harvestman.

The threat sensitive hypothesis predicts that animals modulate the defensive behaviour with the level of threat. Therefore, responses to predator cues may differ from responses to the actual predator in close range. Also, in high threat situations, prey would be expected to use their most dangerous defences. The recluse spider Loxosceles gaucho (Araneae, Sicariidae) is known to prey upon well defended harvestmen such as the laniatorid Mischonyx cuspidatus (Opiliones, Gonyleptidae), which has been reported to use tanathosis, chemical defences, pinching with sharp apophyses on legs, chelicerae and pedipalps. Because of harvestmen’s dependence on chemical stimuli, we tested if M. cuspidatus would change its locomotory behaviour in the presence of chemicals of the recluse spider (low threat situation: spider vs blank vs chemical control; one at a time). Subsequently, we tested harvestmen behaviour in the presence of the spider in close range, a high-threat situation. Finally, we looked at the survival rate of spiders after being pierced by sharp apophyses that M. cuspidatus have on legs IV. The harvestmen only showed defensive behaviours in the high threat situation. Surprisingly, their mostly known defensive behaviours (chemical defence, tanathosis, pinching with chelicerae and pedipalps) were not seen even in the high threat situation. This is the first evidence that these behaviours are not used against a natural predator that has an almost 80% predation success when attacking harvestmen. Pinching with the sharp legs IV apophyses may perforate but do not kill the spiders. We highlight the importance of the traditional descriptive approach with natural predators to understand the specificities of defensive behaviours against different types of predator.

Food consumption in animals is a complex task with multiple steps. Choosing an adequate foraging site is the very first one, and involves not only the presence of prey and predators but also abiotic conditions. Because spiders are usually cannibalistic, conspecifics fall within these two categories in addition to being competitors. Specifically for ambush and cursorial spiders, the type of substrate is also very relevant because spiders often rely on substrate-borne vibrations to find their prey, and distinct substrates propagate vibrations differently. At this point or after contacting the prey, spiders have to decide whether or not to attempt capture. Such a decision involves profitability, prey defenses, and the physiological state of the spider. To capture prey, ambush and cursorial spiders may rely on web sheets, adhesive setae on the tips of the legs, glue-spitting, and venom directly injected from the fangs of the chelicerae. The actual mode of ingestion also varies among species. For almost every step from picking a foraging place to prey consumption, multiple sensory modalities may be used, such as vision, contact chemoreception, olfaction, detection of substrate-borne vibrations, and air displacement. Adequately choosing where to forage, properly detecting, choosing, capturing, and handling prey may have important fitness implications. In this chapter, we summarize the knowledge on these topics with regard to Neotropical cursorial and ambush spiders, detecting gaps and areas better covered within the topics above. Finally, we attempt to suggest promising model species to investigate these different steps of foraging in these animals.

Several arachnid species use chemicals to detect sexual partners. In harvestmen, there are evidences that chemicals may play a role in intraspecific communication. Using the behavior of Discocyrtus prospicuus (Holmberg 1876), whose males expose the penis to females before they engage in mating posture, we tested if males detect females by contact chemoreception (chemicals left on the substrate) and if males detect females by olfaction. First, we exposed males to three experimental groups, where males had to choose between two substrates: female chemicals/blank control; male chemicals/blank control; female/male chemicals. Then, we gave males access to volatiles of males, females, and control simultaneously. We predicted that males would expose the penis when approaching volatiles and chemicals deposited on the substrate by females. We also tested if males spent more time close to the source of female volatiles and on the substrate with female chemicals and if males tapped the substrate with female chemicals for more time than the others. Finally, we put males and females together to observe if males would expose the penis upon touching the female’s cuticle. Most of our predictions were not supported, though males did tap for more time when exposed to female cues instead of male cues and exposed the penis in 70% of the observations when interacting with the female but only after touching her. Our data does not support olfaction as a way to detect females and corroborate the idea that contact chemicals, either on the substrate or on female’s cuticle, play an important role in the detection and recognition of the opposite sex. This is the first evidence in harvestmen that males may react differently to female/male chemicals.

The postembryonic development of Opiliones (Arachnida) includes three phases: larval, nymphal (with four to eight instars), and adult (when molts cease). The present study aimed to describe the postembryonic development of Heteromitobates albiscriptus (Mello-Leitao, 1932) (Gonyleptidae) including both a morphometric study and SEM analysis of two structures present in the tarsus of nymphs and adults: the “tarsal aggregate pores” (TAPs) and the “tarsal perforated organ” (TPO). The nymphal phase includes five stages, which can be easily recognized by morphometric values. In contrast to the pectinate tarsal claws found in legs III–IV of adults (the main synapomorphy of the genus Heteromitobates in the subfamily Goniosomatinae), nymphs bear smooth claws. First nymphs lack TAPs and TPOs. TAPs seem to have a precisely defined position in both prolateral and retrolateral faces of the tarsus. The number of pores in TAPs grows from three or four among second nymphs to around 20 among adults, and measure around 2.15 μm in diameter with no clear difference between ages. An additional field of pores on legs III–IV (“ventral tarsal aggregate pores”, vTAPs) was detected only among adults. The plates at the base and the apex of the TPOs differ from the ones in between. The length of the TPO and its number of plates increase with each molt. However, there is no discernible pattern of growth throughout the postembryonic development when taking into account both the average size of the plates (ranging between ∼7–11 μm) and the ratio of TPO length to tarsus length.

Generalist predators have to deal with prey with sometimes very different morphologies and defensive behaviors. Therefore, such predators are expected to express plasticity in their predation strategy. Here we investigated the predatory behavior of the recluse spider Loxosceles rufipes (Araneae, Sicariidae) when attacking prey with different morphologies and defensive mechanisms. We expected L. rufipes to show different prey capture strategies and variable acceptance towards each prey type. Potential prey species were collected directly from the web or in the surroundings of the web-building site of L. rufipes. We collected and used the following in our experiments: termite workers (Nasutitermes sp.), lepidopteran larvae (Eurema salome), ants (Camponotus sp.) and isopods (Tylidae). We paired these prey with L. rufipes and recorded their behavior in captivity, quantifying acceptance rate, immobilization time and the sequence of behaviors by the predator. The acceptance rate was lower for isopods but not different among other prey. The immobilization time was higher for isopods than for termites and similar for the other pairwise comparisons. The behavioral sequence was similar for all prey except for isopods, which were also bit more often. Our combined results show plasticity in the behavior of L. rufipes and also show it subdues a potentially dangerous prey (ant) and an armored prey (isopod).

The lack of data on the natural history often hampers phylogenetic studies on the evolution of behavior. Herein we provide quantitative field data on foraging and oviposition sites of a Neotropical harvestman belonging to a subfamily with a published phylogeny, Goniosomatinae. Heteromitobates discolor rests during the day on granitic boulders on rivers, laying eggs in sheltered and darker areas. The female guard the eggs and aggressively respond to approaching conspecific females. In the absence of the female, ants, conspecifics and reduviids may predate the eggs. Egg-guarding females are known not to leave the clutch, but males and non-guarding females may leave the granitic boulders and forage on the vegetation at night, close to the river margins and with no difference in the sites explored between males and females. The general features described in H. discolor are similar to what has been reported for other species in this subfamily, suggesting an evolutionary conservatism within the group.

Males of Iporangaia pustulosa (Arachnida: Opiliones) have a sexually dimorphic metatarsus IV, which is thicker and with more glandular pores in males. Here we tested the hypothesis that this glandular area is used by males to leave chemicals in the environment, predicting that the animals would rub the metatarsus IV against the substrate. We have made recordings both in the field and in the laboratory, in several distinct contexts during the day and at night, comprising 67 hours of observations. We also experimentally tested the reaction of both sexes to a filter paper rubbed on the metatarsus gland, with adequate controls. We report and describe for the first time that the metatarsal gland of I. pustulosa is used to leave chemicals on the substrate by rubbing or touching it against the substrate. We also provide evidence that males can control the release of secretions of the metatarsal gland IV.

O estudo da personalidade em animais é relativamente recente. Até pouco tempo atrás, na Ecologia Comportamental, as diferenças individuais consistentes eram consideradas ruído em torno à estratégia ótima. Na Psicologia, evitava-se falar de personalidade em animais por temor à antropomorfização. As pesquisas têm mostrado que diferenças de personalidade se manifestam em inúmeras espécies, como formas diferentes de lidar com as pressões seletivas, com consequências para a aptidão, sendo assim uma variável muito importante em ambientes que mudam. Temperamento, Personalidade, Síndrome Comportamental ou Estilo de Enfrentamento são nomes que têm sido usados para se referir ao fenômeno dos animais exibirem diferenças interindividuais consistentes ao longo do tempo e das situações. Porém, esses termos não são exatamente sinônimos. Ao longo desta tese, primeiro revisei o percurso histórico do estudo da personalidade em animais a partir das perspectivas da Psicologia e da Ecologia Comportamental, buscando os pontos em comum e as possibilidades de unificação das duas escolas. No segundo capítulo, avaliei se houve diferenças de personalidade, mediante a codificação do comportamento, em doze macacos-prego selvagens observados dentre 0 e 3 anos de idade. Para tanto, verificamos, usando uma análise de repetibilidade, se as diferenças individuais foram consistentes ao longo de 10 pontos de desenvolvimento, para conferir se as características de personalidade estavam já presentes logo ao nascimento ou se sofreram modificações ao longo x do desenvolvimento. Mediante análise dos componentes principais obtivemos quatro traços de personalidade: Sociabilidade-Atratividade, Ansiedade, Apertura e Atividade, achando um efeito do desenvolvimento em Sociabilidade (que diminuiu com a idade) e Ansiedade (que aumentou). Contudo, não foi possível verificar consistência intra-individual nos traços ao longo do tempo analisado. Em consonância com a literatura de personalidade e temperamento em humanos, os resultados indicaram que a estrutura da personalidade não está ainda estabelecida no começo do desenvolvimento The study of animal personality is relatively new. Until recently, in Behavioral Ecology, consistent individual differences were considered \'noise\' around an optimal strategy. In Psychology, animal personality was avoided as a consequence of the fear of anthropomorphization. Research has shown that personality differences are manifest in many species, evident from different forms of dealing with selective pressures, with consequences for fitness, thus, being an important variable in changing environments. Temperament, Personality, Behavioral Syndrome, or Coping Style are names that have been used to refer to the phenomenon of animals exhibiting \"consistent interindividual differences over time and across situations. However, these terms are not exactly synonyms. Throughout this thesis, I first reviewed the historical course of the study of personality in animals, from the perspectives of Psychology and Behavioral Ecology, looking for points in common and possibilities of unification of the two schools. In the second chapter, I evaluated, through behavioral coding, if there were personality differences in twelve young wild capuchin monkeys, between 0 and 3 years-old. Then, we verified using a repeatability analysis whether the individual differences were consistent across 10 developmental points, to find out if the personality traits were already present at birth or if they were acquired during development. By means of principal component analysis, we obtained four personality traits: Sociability-Attractiveness, Anxiety, Openness and Activity, finding an effect of development on Sociability (which decreased xii with age) and Anxiety (which increased). However, it was not possible to verify intra-individual consistency in the traits throughout the analyzed time. Consistent with the literature on personality and temperament in humans, the results indicated that personality structure is not yet established at the beginning of development

As interações entre presas e predadores possuem um papel fundamental na evolução das espécies. Para as presas em particular, uma falha na interação com um predador pode representar a morte e, consequentemente, os genes desta presa provavelmente não estariam representados em gerações futuras. Devido à essa forte pressão seletiva imposta pelos predadores, as presas evoluíram diversas estratégias defensivas. As defesas podem atuar dificultando que os predadores detectem a presas (e.g. anacorese e a cripsia) ou após os predadores estarem muito próximos ou mesmo terem atacado às presas. Tais defesas incluem a retaliação (tanto mecânica como química) e o comportamento de tanatose. Todas estas defesas mencionadas anteriormente foram propostas para opiliões. Portanto, opiliões são potencialmente bons modelos para estudar a evolução das estratégias defensivas. Nesta tese, adotamos o opilião Mischonyx cuspidatus como modelo experimental. No capítulo 1, testamos se M. cuspidatus são consistentes nos níveis de ousadia (propensão em se exporem ao risco, medido neste trabalho pelo tempo em tanatose e em freezing) e se a ousadia é influenciada pelo jejum. Encontramos que a ousadia é consistente, mas não detectamos efeitos do jejum. No capítulo 2, testamos se existem diferenças sexuais nas defesas de M. cuspidatus. Considerando o dimorfismo sexual presente nesses animais (machos possuem estruturas pontiagudas no último par de pernas que são utilizadas retaliação, mas as fêmeas não), testamos as seguintes predições i) machos dependeriam mais de defesas mecânicas do que fêmeas e ii) fêmeas dependeriam de outras formas de defesa, como defesas químicas e defesas passivas. Encontramos que os machos respondem com retaliação mecânica mais frequentemente do que as fêmeas e as fêmeas realizam tanatose mais frequentemente do que os machos. Não encontramos diferenças na frequência de uso de defesas químicas. No capítulo 3, testamos se há diferenças entre os períodos diurno e noturno na realização do 466 comportamento de tanatose em fêmeas de M. cuspidatus. Prevemos que as fêmeas realizariam tanatose mais frequentemente durante o dia. Assim, poderiam potencialmente evitar que fossem detectadas por predadores visuais. Conforme esperado, encontramos que fêmeas realizam tanatose mais frequentemente durante o dia do que à noite. Os resultados que obtivemos nesta tese contribuem para o conhecimento da evolução das estratégias defensivas, bem como fornece resultados que podem ser um passo inicial para o entendimento da consistência comportamental em opiliões e suas implicações para este táxon Prey-predator interactions play a major role in species evolution. Failing in an interaction with a predator may represent death for prey. Consequently, prey´s genes would not be represented in the future generations. In response to the strong selective pressures imposed by predators, prey have evolved several defensive strategies. Prey defenses may hinder detection by predators (e.g. anachoresis and crypsis). Other defensive strategies may play a role after predators are in close-range or already attacked the prey, for example retaliation (both mechanical and chemical) and thanatosis. All the aforementioned defenses have been described to harvestmen. Thus, harvestmen are potentially good models to study the evolution of defensive strategies. In this thesis, we used the harvestman Mischonyx cuspidatus as our experimental model. In the first chapter, we tested if individuals of M. cuspidatus are consistent in boldness levels (propensity to take risks, herein measured by the time spent performing thanatosis and freezing). We also tested if boldness is influenced by starvation. We found that M. cuspidatus behaved consistently regarding boldness, but we did not find any effect of starvation. In the second chapter, we looked for sexual differences in defensive behavior of M. cuspidatus. Taking into account the sexually dimorphic characters of this species (males have sharp structures in the fourth pair of legs, but females do not) we tested the following predictions i) males would rely more upon mechanical defenses than females and ii) females would rely more on other kinds of defenses, such as chemical and passive defenses than males. We found that males performed mechanical retaliation more often than females and females performed thanatosis more often than males. We did not find any sexual difference with respect to chemical defense. In the third chapter we tested if there are differences in the frequency of thanatosis performed by females of M. cuspidatus between the periods of night and day. By performing thanatosis more often during the day, females of M. cuspidatus might be able to avoid being detected by visual predators. As expected, females performed thanatosis more often during the day than at night. Taken together, our results contribute to the understanding of the evolution of defensive strategies, as well as provide results that might be a first step to address new questions about behavioral consistency in harvestmen and their implications in this group

Capítulo 1: Dimorfismo sexual em aberturas glandulares das pernas de três subfamílias de Gonyleptidae (Arachnida: Opiliones: Laniatores) Glândulas sexualmente dimórficas podem não apenas ser informativas taxonomicamente como podem ainda trazer informações importantes sobre a comunicação entre os sexos. Nesse trabalho estudamos a evolução de aberturas glandulares sexualmente dimórficas nas pernas I (basitarso) e pernas IV (presença/ausência de intumescimento e quantidade de poros no metatarso) em uma linhagem monofilética pertencente ao clado K92 composta pelas subfamílias Progonyleptoidellinae, Sodreaninae e parte de Gonyleptinae. Utilizamos Microscopia Eletrônica de Varredura para descrever as aberturas glandulares e compará-las entre exemplares das três subfamílias analisadas. Encontramos aberturas glandulares sexualmente dimórficas na perna I de uma espécie em Progonyleptoidellinae (de seis analisadas), de uma espécie de Gonyleptinae (de quatro analisadas) e em três espécies de Sodreaninae (de seis analisadas). Com relação às aberturas da perna IV, todos os machos estudados possuem maiores quantidades de poros na perna IV do que as fêmeas exceto uma espécie de Gonyleptinae e uma de Sodreaninae. Os machos de todas as espécies estudadas possuem o astrágalo das pernas IV intumescidos, exceto uma espécie em Progonyleptoidellinae. Portanto as aberturas glandulares sexualmente dimórficas na perna I nos machos mostram-se variáveis na linhagem interna ao clado K92 estudada nesse trabalho. Por outro lado, o intumescimento e quantidade de poros na perna IV dos machos parecem ser conservados na linhagem estudada. Capítulo 2: Morfologia interna das glândulas sexualmente dimórficas de Progonyleptoidellus striatus Roewer 1913 (Opiliones: Laniatores: Gonyleptidae) Apesar de diversos trabalhos sugerirem a presença de glândulas sexualmente dimórficas nas quatro subordens de Opiliones, há poucos trabalhos que investigam essas glândulas histologicamente. Morfologia interna fornece importantes informações sobre o funcionamento e composição das secreções. Para isso, utilizamos técnicas histológicas e ultraestruturais para estudar a morfologia interna de duas glândulas de Progonyleptoidellus striatus (Progonyleptoidellinae): uma no basitarso das pernas I e a outra no metatarso das pernas IV. Adicionalmente, investigamos a composição histológica e ultraestrutural da área glandular no metatarso IV. Encontramos diferenças entre os sexos, tanto na morfologia interna como externa: machos possuem intumescimentos das áreas glandulares e internamente possuem células glandulares ausentes nas fêmeas. Encontramos abundantes retículos endoplasmáticos rugosos, lisos e mitocôndrias, além de Complexo de Golgi bem desenvolvido na perna I dos machos. Diferentemente da perna I, não encontramos retículos endoplasmáticos lisos na perna IV, o que pode implicar em composições de produtos secretórios diferentes para as pernas. Nas duas pernas encontramos abundância de vesículas secretoras Sexually dimorphic glands may not only be taxonomically informative but can also bring important information about communication between the sexes. In this work we studied the evolution of sexually dimorphic glandular openings on legs I (basitarsus) and legs IV (presence/absence of swollen area and quantity of pores on metatarsus) in a monophyletic lineage belonging to clade K92 that include the subfamilies Progonyleptoidellinae, Sodreaninae and part of Gonyleptinae. We used Scanning Electron Microscopy to describe the glandular openings and compare them among specimens of the three subfamilies analyzed. We found sexually dimorphic glandular openings on legs I of one species in Progonyleptoidellinae (out of six analyzed), one specie of Gonyleptinae (out of four analyzed) and three species of Sodreaninae (out of six analyzed). Concerning glandular openings on legs IV, all males studied have more pores than females except for one species in Gonyleptinae and another in Sodreaninae. Males of all species studied have a swollen astragalus IV, except for one species in Progonyleptoidellinae. Therefore, the glandular openings in leg I in males are shown to be variable for the internal lineage to the Clade K92 studied in this work. On the other hand, the swollen area and the greater quantity of pores on legs IV of males seem to be conserved in the lineage studied. Although several studies suggest the presence of sexually dimorphic glands in the four suborders of Opiliones, there are few studies that investigate these glands histologically. Internal morphology provides important information about the functioning of the glands and the composition of secretions. Here we used histological and ultrastructural techniques to study the internal morphology of two glands of Progonyleptoidellus striatus (Progonyleptoidellinae): one in the basitarsus of legs I and another in the metatarsus of legs IV. We found differences between the sexes, both in internal and external morphology, in which males have swollen of the glandular areas and internally the males have glandular cells, absent in females. We found abundant rough, smooth endoplasmic reticulum and mitochondria, as well as Golgi Complex well developed in legs I of males. In contrast, we did not find smooth endoplasmic reticulum in legs IV, which may result in different compositions of glandular products in each leg. In both legs we found abundance great number of secretory vesicles

O comportamento gregário é comum em algumas espécies de Opiliones. No período noturno indivíduos gregários saem do abrigo para forragear e ao amanhecer retornam ao ambiente de repouso. Químicos de coespecíficos têm sido discutidos como mediadores do retorno ao abrigo nos indivíduos da subordem Eupnoi, mas até o momento nada se sabe sobre as pistas que os indivíduos da subordem Laniatores utilizam para retornar ao abrigo. Recentemente, uma grande diversidade de receptores olfativos foi descrita nessa subordem e o uso do olfato à curta distância (< 2 cm) já foi documentado em agregados. Testamos as hipóteses do uso do olfato à longa distância (50 cm) na atração por (1) químicos voláteis emitidos diretamente de coespecíficos e (2) químicos voláteis próprios e de coespecíficos deixados previamente nos abrigos. Utilizamos um olfatômetro em formato de Y. Usamos como estímulos fêmeas e machos coespecíficos para (1) e abrigos dos sujeitos experimentais, de coespecífico do mesmo sexo e controle para (2). Para (1) machos de M. cuspidatus escolheram significativamente o lado do olfatômetro com machos coespecíficos. Fêmeas de M. cuspidatus não mostraram preferência por nenhum dos estímulos. Para (2) machos e fêmeas de M. cuspidatus não escolheram significativamente nenhum estímulo do olfatômetro. Esses resultados nos mostram que talvez machos de M. cuspidatus tenham maior sensibilidade olfativa comparada ao das fêmeas coespecíficas. Os machos de M. cuspidatus não mostraram preferência por fêmeas coespecíficas talvez porque no momento do experimento as fêmeas não estivessem liberando voláteis. Nossos resultados também mostram que indivíduos de M. cuspidatus não são atraídos à distância por químicos deixados no abrigo, talvez porque esses químicos sejam pouco voláteis ou porque os indivíduos detectaram o odor, mas optaram por não escolher o braço com odor próprio e nem de coespecífico. O olfato parece ser mais importante do que anteriormente mencionado em Opiliones Gregarious behavior is common in some species of Opiliones. At night, gregarious individuals leave their shelter to forage and at dawn return to the original spot. Chemicals of conspecifics have been suggested to be mediators of homing in individuals of the suborder Eupnoi, but so far nothing is known about the cues used in the suborder Laniatores. Recently, a large diversity of olfactory receptors has been described in this suborder and the use of olfaction at close range (

A interação envolvendo artrópodes e plantas é uma das mais antigas e importantes interações ecológicas da natureza. As estratégias adotadas pelos artrópodes para reconhecer a sua planta hospedeira são bastante variáveis e pistas químicas desempenham um papel importante nessa relação. Opiliões (Arachnida, Opiliones) são altamente dependentes de quimiorrecepção para encontrar recursos e são particularmente dependentes de umidade. O opilião Jussara sp. possui uma clara preferencia por repousar em Psychotria suterella (Rubiacea), uma planta com uma arquitetura foliar complexa com folhas sobrepostas. Nada se sabe sobre como o opilião chega a ela ou quais as vantagens que ela proporciona ao opilião. Neste trabalho verificamos quais pistas são utilizadas pelo opilião para encontrar P. suterella e ainda quais os possíveis benefícios para o opilião nessa interação. Para verificação de como o opilião encontra a planta, partimos de duas hipóteses não excludentes: 1 - pistas de co-específicos são utilizadas; 2 - pistas químicas provenientes da planta são utilizadas. Para a primeira hipótese, montamos um experimento em campo com um grupo de plantas de P. suterella contendo químicos de Jussara sp. e um grupo sem químicos de Jussara sp. no caule principal e comparamos como os tratamentos afetaram o número de opiliões nas plantas de cada grupo. Para a hipótese da utilização de químicos da planta, realizamos dois experimentos visando verificar a preferência do opilião pelos voláteis de P. suterella. No primeiro utilizamos extratos de químicos voláteis de P. suterella e de mais dois controles em uma arena triangular com os 3 estímulos oferecidos simultâneamente. No segundo experimento utilizamos a própria planta P. suterella, pareada com uma planta controle ou um controle branco em um olfatômetro em Y. Para verificar os benefícios da associação também partimos de duas hipóteses não exludentes relacionadas a possíveis diferenças microclimáticas geradas por P. suterella em relação às demais plantas do local: 1- as folhas de P. suterella oferecem maior umidade do que outras plantas; 2 - as folhas de P. suterella fornecem um microambiente mais sombreado do que o de outras plantas. Para primeira hipótese, quantificamos o fluxo de transpiração foliar de P. suterella e mais 3 espécies de plantas do habitat do opilião. Já para segunda hipótese, quantificamos o bloqueio à passagem de luz através das folhas em P. suterella e das mesmas 3 espécies do teste anterior. Não obtivemos evidências claras de que os opiliões Jussara sp. utilizem químicos de co-específicos para selecionar P. suterella. Pistas químicas olfativas não parecem ser utilizadas de forma isolada para atrair Jussara sp., mas nossos resultados sugerem que talvez em combinação com outras plantas o odor possa ser utilizado. Em relação aos benefícios fornecidos por P. suterella, nós não encontramos evidências de que a transpiração foliar seja importante na interação. Contudo, o maior bloqueio de passagem de luz através da folha em comparação às demais espécies aliado a arquitetura foliar de P. suterella pode gerar microclimas benéficos para os opiliões que geralmente são altamente sensíveis a umidade Interactions between arthropods and plants are among the oldest and important ecological interactions in nature. Strategies adopted by arthropods to recognize its host plant are very variable and volatiles often mediate the encounter between both parts. Harvestmen (Arachnida, Opiliones) are arachnids highly dependent on chemicals to find resources and particularly dependent on high humidity. The harvestman Jussara sp. clearly prefers to rest on Psychotria suterella (Rubiacea), a plant with a complex architecture with overlapping leaves. We know nothing about how the harvestmen find the plant or the advantages that such plant offers to the harvestmen. In this paper we investigate the cues used to find P. suterella by these harvestmen and how the harvestman benefits from this interaction. To test how the harvestman finds the plant, we raised two non-excludent hypotheses: 1 - conspecific chemicals are used; 2 - plant chemicals are used. For the first hypothesis, we conducted a field experiment with a group of P. suterella with chemicals of Jussara sp. and another without chemicals of of Jussara sp. on the main stem and compared how that affected the number of harvestmen on them. To test the importance of plant chemicals, we conducted two experiments to test the use of volatiles of P. suterella. We first used volatile extracts of P. suterella and two other plants in a triangular arena with the three stimuli offered simultaneously. In the second experiment, we used actual individuals of P. suterella paired with either a control plant of a white control in a Y maze. To investigate how Jussara sp. benefits by choosing P. suterella, we raised two non-excludent hypotheses related to microclimatic conditions potentially offered by P. suterella in comparison to other plants of the area: 1 - The leaves of P. suterella offer higher humidity than those of other plants; 2 - The leaves of P. suterella provide a better shadow that that of other plants. For the first hypothesis, we quantified the transpiration rate of the leaves of four local species including P. suterella. For the second hypothesis, we quantified how much light passes through the leaves of these four species. We did not find clear evidences that Jussara sp. uses conspecific chemicals to select P. suterella. Plant volatiles do not seem to be used at least isolated to attract Jussara sp., but our results suggest that maybe in combination with other plants such volatiles may be used. As for the benefits provided by P. suterella, we did not find evidence that the transpiration rate is important. However, the leaves of P. suterella do provide better shadows in comparison with the other plants, which in combination with the complex architecture with overlapping leaves may provide better microclimatic conditions to these very humidity sensitive animals

O conhecimento sobre cestódeos parasitas de aves na região Neotropical é bastante escasso, sendo baseado em materiais coletados nos séculos XVII e XVIII. No Brasil, a diversidade reportada para estes helmintos é baixa e não é compatível com a grande diversidade de aves registrada no país. Buscou-se inventariar a riqueza e a diversidade de cestódeos parasitas de aves da família Thamnophilidae no sul da região Amazônica brasileira, nos seus principais interflúvios e centros de endemismo, afim de caracterizar a fauna de cestódeos e verificar sua distribuição em relação à observada para os hospedeiros. Ao todo foram amostrados 487 tamnofilídeos pertencentes a 81 táxons, distribuídos nos principais interflúvios e centros de endemismo ao sul da Amazônia brasileira, no levantamento mais extenso do ponto de vista geográfico e temporal já realizado para Aves na América do Sul. As técnicas tradicionais de coleta e amostragem de cestódeos foram aperfeiçoadas levando-se em conta as especificidades de clima e logística características da região amazônica, resultando na coleta de aproximadamente 1.500 indivíduos de cestódeos das famílias Dilepididae, Hymenolepididae, Paruterinidae e Metadilepididae, pertencentes à Ordem Cyclophyllidea. Os parasitas encontrados podem representar 83 novas espécies, além de novos gêneros em todas as famílias de cestódeos amostradas. As chaves de identificação disponíveis para estes grupos não são suficientes para permitir a identificação dos táxons encontrados parasitando os Thamnophilidae. A consideração da especificidade parasita-hospedeiro para a identificação dos helmintos obtidos é um impedimento no refino taxonômico para este grupo, pois não há cestódeos descritos formalmente parasitando a família Thamnophilidae e é necessária uma ampla revisão das chaves de identificação disponíveis. A técnica de Microscopia Eletrônica de Varredura foi utilizada pela primeira vez para caracterizar os táxons da família Dilepididae, mostrando-se informativa na distinção dos morfotipos. A relação de especificidade parasita-hospedeiro não foi verificada na maioria dos casos, sendo o mais frequente uma relação ecológica entre hábitos alimentares dos hospedeiros e as diferentes famílias de parasitas. Verificou-se que os grandes rios amazônicos não atuam como barreiras para os parasitas, sendo encontrados os mesmos morfotipos em táxons de aves distintos e separados pelos rios. A influência das relações ecológicas entre os hospedeiros parece ser um fator mais relevante na distribuição dos parasitas, pois diferentes populações de aves em um mesmo interflúvio apresentaram uma comunidade de cestódeos distinta. A diversidade de cestódeos ainda é extremamente subestimada na região Neotropical, e este impedimento taxonômico dificulta a realização de análises mais aprofundadas, sendo fundamental que a taxonomia esteja mais bem resolvida para os membros desta Classe. Este estudo foi pioneiro na caracterização dos cestódeos parasitas de Thamnophilidae e abre uma nova e promissora linha de pesquisa nesta área em nada explorada The information about tapeworms inhabiting Neotropical birds is scarce, based on samples collected from XVII and XVIII centuries. In Brazil, these helminths diversity is low and not consistent with the high birds diversity. In this study, we examined the abundance and diversity of tapeworms inhabiting antbirds of family Thamnophilidae in southern Brazilian Amazon, in areas of endemism between the major rivers, to report the tapeworm fauna and its distribution regarding the hosts distribution. As result, we sampled 487 antbirds of 81 taxa, along the major interfluvial areas and areas of endemism is southern Brazilian Amazon, in the more extensive survey of parasites in South American birds. The traditional protocols for sampling cestodes were improved based on climate conditions and organization of Amazon area, resulting in a sample about to 1500 Cyclophyllidean cestodes belonging to families Dilepididae, Hymenolepididae, Paruterinidae e Metadilepididae. The specimens collected may represent 83 new species, besides new genera for all the families. The keys for Cyclophyllideans tapeworms are not useful for Thamnophilidae parasites, demanding revision for considering the host-parasite specificity and no parasites described inhabiting antbirds. For the first time Scanning Electronic Microscopy is used to discriminate Dilepididae species successfully. The host-parasite specificity for Thamnophilidean tapeworms is low and the parasite diversity is more influenced by the host foraging habits. The major Amazonic rivers are not physiographic barriers for cestodes distribution and the same parasites morphotype were reported in different host from both sides of a river. The ecological relationships between the hosts in a same area can influence their parasite diversity, with distinct birds populations in the same area parasited by different cestodes. The Neotropical tapeworm diversity is underestimated and this interferes in broader studies for host-parasites relationships, demanding taxonomic revision for these parasites class. This is the first survey for Thamnophilidae tapeworms and suggests a new and promising unknown line of research

Opiliões (Arachnida, Opiliones) são especialmente dependentes da quimiorecepção de contato, além de serem muito dependentes de altos níveis de umidade e de temperaturas amenas. No entanto, o conhecimento acerca das estruturas que detectam esses estímulos é muito limitado em opiliões, quando comparado ao que se sabe sobre outros aracnídeos. Além disso, são raros os estudos investigando a morfologia interna dessas estruturas - um tipo de informação essencial para a determinação de suas funções- e o potencial das sensilla para a sistemática. Neste trabalho, nós investigamos as sensilla tarsais de todos os pares de perna do opilião Heteromitobates discolor (Laniatores, Gonyleptidae), buscando refinar o conhecimento das sensilla quimioreceptoras e investigar a localização dos receptores de temperatura e umidade, através de microscopia eletrônica de varredura (MEV) e transmissão. Para determinar a abrangência dos resultados obtidos com H. discolor e testar o uso de algumas estruturas para a sistemática do grupo, realizamos uma amostragem com MEV em espécies das quatro subordens de Opiliones (Cyphophthalmi, Eupnoi, Dyspnoi e Laniatores), com foco em Laniatores (subordem com 2/3 das espécies do grupo). Na primeira parte, fornecemos a primeira evidência morfológica de receptores olfativos em Laniatores (em H. discolor), mostrando que há abundância e diversidade de sensilla olfativas. Além disso, fornecemos evidência (com MEV) de cerdas olfativas em 17 famílias de Laniatores, o que sugere que a olfação é mais importante para os Laniatores do que previamente considerado. Na segunda parte, fornecemos a primeira evidência morfológica de detectores de umidade e temperatura em opiliões (em H. discolor), discutindo os mecanismos de funcionamento dessas estruturas e uma associação morfológica entre elas, localizada na parte mais distal das pernas I e II. Por fim, mostramos que essas sensilla candidatas a detectores de umidade e temperatura são extremamente conservadas em Laniatores (28 famílias), e que existem estruturas comparáveis em espécies de Cyphophtalmi, Eupnoi e Dyspnoi. Com uma análise de reconstrução de estado ancestral em uma filogenia de Opiliones compilada da literatura, mostramos que as variações na morfologia externa dessas estruturas fornecem informação filogenética em vários níveis de relacionamento em Opiliones. Esse estudo contribui para o conhecimento de aspectos básicos da anatomia celular de sensilla em Opiliones, refinando o conhecimento sobre a função das sensilla tarsais e forncendo uma base para fomentar o uso de sensilla para a sistemática do grupo Harvestmen (Arachnida, Opiliones) are especially dependent on contact chemoreception and are dependent on high humidity levels and amenable temperatures. However, knowledge on the sensory structures (sensilla) that detect such stimuli is limited in harvestmen when compared with other arachnid orders. Besides, there are few studies investigating the internal morphology of these structures -which is important for inferring function - and the potential of sensilla for systematics. To refine the knowledge on chemoreceptive sensilla and investigate the identity of hygro- and thermoreceptors, we investigated the tarsal sensilla of all leg pairs of the species Heteromitobates discolor (Laniatores, Gonyleptidae), using scanning electron microscopy (SEM) and transmission electron microscopy. To determine scope of the results obtained with H. discolor and to test the use of some structures for systematics, we surveyed (with SEM) species in all four suborders of Opiliones (Cyphophthalmi, Eupnoi, Dyspnoi and Laniatores), with focus in Laniatores (suborder with 2/3 of harvestmen species). In the first part, we provide the first morphological evidence of olfactory receptors in a species of Laniatores (H. discolor), showing that olfactory sensilla are abundant and diverse. Also, we show evidence (with SEM) of olfactory sensilla on 17 families of Laniatores, which suggests that olfaction is more important for Laniatores than previously considered. In the second part, we provide the first morphological evidence of hygro- and thermoreceptors in harvestmen (in H. discolor) discussing the functioning mechanisms of these structures and their morphological inter-association on the distal-most part of leg pairs I and II. Finally, we show that these putative hygro- thermoreceptive sensilla are widespread in species of Laniatores (28 families) and that comparable structures occur on species of the suborders Cyphophthalmi, Eupnoi and Dyspnoi. With an ancestral state reconstruction on a compiled phylogeny of Opiliones, we show that morphological variations on these sensilla are informative on several levels of phylogenetic relationships in Opiliones. This study constributes for the basic knowledge on the cellular anatomy of Opiliones sensilla, refining the function of tarsal sensillar types and providing a base for their use in systematics

As mudanças climáticas influenciarão as médias de temperaturas ambientais e a frequência de eventos extremos com consequências ainda desconhecidas para a fauna. Conhecer os limites fisiológicos dos organismos ao aumento de temperatura pode ajudar a ampliar os marcos conceituais dos impactos regionais das mudanças climáticas sobre a fauna. Este trabalho abordou como a diversidade fisiológica representada pela tolerância termal da fauna de artrópodes terrestres estava relacionada entre os biomas do Brasil, em uma perspectiva macrofisiológica (ampla escala espacial). Especificamente, coletou-se a temperatura crítica máxima (TCMax) de espécimens de artrópodes das Classes Arachnida, Chilopoda, Dipoploda, Entognatha (Collembola), Insecta e Malacostraca (Oniscidea) nos biomas de Mata Atlântica (strictu sensu), Mata Atlântica de Altitude, Cerrado, Amazônia e Caatinga. Os artrópodes foram utilizados como modelo de estudo por permitirem uma associação mais clara com a teoria disponível sobre limites fisiológicos e o ambiente físico. Assim, foram investigados padrões de variação da TCMax entre e dentro dos biomas, considerando ou não a categoria sistemática. Igualmente, foi avaliada a relação da TCMax da fauna de artrópodes com variáveis bioclimáticas representantes do ambiente termal nos biomas. No aspecto de margem de segurança termal, avaliou-se a potencial tolerância ao aquecimento da fauna de artrópodes nos biomas. Os resultados mostraram que existe ampla diversidade fisiológica da fauna de artrópodes, com padrões atrelados aos biomas do Brasil. A relação do padrão de tolerância termal dos espécimens de artrópodes com o bioma é complexa, havendo nuances de variação dentro e entre os biomas. Houve grande proporção de fauna termotolerante no bioma da Caatinga e menor proporção de fauna termotolerante na Mata Atlântica. Quanto às margens de segurança termal, os dados de tolerância ao aquecimento sugerem que não há grande susceptibilidade ao aquecimento da fauna de artrópodes nos biomas do Brasil. Este trabalho contribui para ampliar a discussão dos possíveis impactos das mudanças climáticas em seus aspectos regionais, tendo em vista a diversidade fisiológica da fauna de artrópodes terrestres nos biomas brasileiros. Igualmente, os dados podem servir como uma importante base para a incorporação em avaliações globais da vulnerabilidade dos ectotérmicos frente às mudanças do clima Climate change will affect the average environmental temperatures and the frequency of extreme events with still unknown consequences for wildlife. To understand the physiological limits of organisms in relation to the increase in environmental temperature can help extend the conceptual frameworks of climate change regional impacts on wildlife. This work discussed how the physiological diversity represented by the thermal tolerance of terrestrial arthropod fauna was related among biomes of Brazil in a macrophysiological perspective (large spatial scale). It was collected critical thermal maximum (CTMax) of specimens of the class Arachnida, Chilopoda, Dipoploda, Entognatha (Collembola), Insecta and Malacostraca (Oniscidea) in the biomes of the Atlantic Forest (strictu sensu), Atlantic Forest Highland, Cerrado (Brazilian Savanna), Amazonia and Caatinga. The arthropods were used as model to allow a better association with the available theory of physiological limits and the physical environment. Thus, it was investigated variation in patterns of CTMax between and within biomes considering or not the systematic category. It was also evaluated the relationship of CTMax of the arthropod fauna with bioclimatic variables as representative of the thermal environment in the biomes. In terms of thermal safety margin, it was evaluated potential warming tolerance of the arthropod fauna in the biomes. The results showed that there is a broad physiological diversity of arthropod fauna with patterns linked to brazilian biomes. The ratio between thermal tolerance patterns of arthropod specimens and the biome is complex, there were varying nuances within and between biomes. There is a large proportion of thermotolerant fauna in the Caatinga biome and a lesser proportion of thermotolerant fauna in the Atlantic Forest. The warming tolerance data suggest that the susceptibility to heat of the arthropod fauna in brazilian biomes is small. This work will help to expand discussions of potential impacts of climate change regional aspects considering the view of the physiological diversity of terrestrial arthropod fauna in the brazilian biomes. In addition, the data can be as an important basis for incorporation into global vulnerability assessments on terrestrial ectotherms in view of climate change

Muitas formas de cuidado parental aumentam a sobrevivência da prole, ao custo de uma diminuição na capacidade dos pais em investir em proles futuras. Espera-se, portanto, que indivíduos parentais adotem estratégias de cuidado que lhes permitam balancear benefícios imediatos e custos futuros, otimizando seu sucesso reprodutivo total. Nesta dissertação, testamos um potencial custo e um benefício do cuidado maternal do opilião Neosadocus maximus, espécie cujas fêmeas desertam suas desovas periodicamente e acrescentam ovos a elas por um período de até duas semanas características únicas entre opiliões com cuidado maternal. Exploramos também quais fatores permitem a deserção temporária das fêmeas de N. maximus. Mais especificamente, testamos as seguintes hipóteses: (1) o cuidado maternal protege os ovos contra predadores, que são mais comuns no período noturno; (2) o cuidado maternal diminui a fecundidade imediata da fêmea; (3) a camada de muco que as fêmeas depositam sobre seus ovos mantém a prole protegida nos momentos de deserção temporária. Nossos resultados apontam que o cuidado maternal é uma proteção eficaz contra o ataque de predadores, sem afetar negativamente a fecundidade imediata das fêmeas. Demonstramos também que a camada de muco mantém os ovos relativamente protegidos na ausência da fêmea guardiã, permitindo que estas se ausentem de suas desovas sem deixar a prole totalmente vulnerável. Sendo assim, a camada de muco que recobre os ovos de N. maximus permite que estas fêmeas minimizem os custos fisiológicos associados ao cuidado maternal sem, entretanto, aumentar os custos da deserção em termos de redução de prole provocada por predação. Many forms of parental care increase offspring survival, at the cost of a decrease in the parental individuals capacity to invest in a future brood. It is expected, therefore, that parental individuals adopt parental strategies that allow them to balance immediate benefits and future costs, optimizing their total reproductive success. In this thesis, we aimed to identify costs and benefits of maternal care in the harvestman Neosadocus maximus, whose females periodically desert their broods and add eggs to their clutches for up to two weeks unique characteristics among harvestmen with maternal care. We also explored which factors allow N. maximus females to desert their clutches frequently. We tested the following hypothesis: (1) maternal care protects the eggs against predators, especially at night; (2) maternal care decreases the current female fecundity; (3) the mucus coat covering the eggs protects them against predators even in the absence of the guarding female. Our results indicate that maternal care is an effective protection against egg predators, and does not decrease current female fecundity. We also demonstrated that the mucus coat provides effective protection to the eggs in the absence of the guarding female, allowing them to abandon periodically their clutches without leaving the offspring completely vulnerable to predators. The mucus coat covering N. maximus eggs allows these females to minimize the physiological costs of caring, with no severe increase in the cost of deserting in terms of brood reduction by predation.

O estudo da divisão de tarefas e de sua participação na manutenção das condições adequadas à sobrevivência de insetos sociais têm recebido importantes contribuições ao longo dos anos. O elaborado grau de organização de muitos insetos sociais e a presença de polimorfismo em muitos grupos com formação de castas, tornam a investigação da divisão de tarefas ainda mais interessante. Essa dissertação apresenta os resultados de três experimentos com colônias de formigas cortadeiras Atta sexdens rubropilosa: 1) Foi verificado se há trofalaxia de água marcada entre companheiras de ninho de A. sexdens rubropilosa em colônias em condição de estresse hídrico. Os resultados encontrados mostraram que a trofalaxia é um evento que ocorre com freqüência em colônias em estresse hídrico. 2) Foi testado a divisão de tarefas associado a dinâmica dos tamanhos envolvidos na atividade de coleta de água as distâncias de 0 metro, 1 metro e 10 metros para colônias em condição de estresse hídrico. Encontramos que após um tempo a coleta de água foi associada a uma casta especializada, de menor porte, ou seja, no início da atividade com formigas de maior porte e com o tempo formigas menores passaram a realizar a tarefa, fato que aqui chamamos de \"Efeito do Tempo\". Este efeito temporal de especialização variou de acordo com a distância a qual a fonte de água foi colocada. 3) Uso do sistema de pontes com fenda de 1,5 cm. Foi elaborado um sistema modelo de ponte com fenda e sem fenda e testado a separação física de tamanhos e a possibilidade de aprendizado da existência da fenda. Os resultados indicam que ocorre uma separação física das formigas maiores. Os dados preliminares sugerem o aprendizado da existência da fenda. Assim este trabalho buscou oferecer uma contribuição as inúmeras conexões que envolvem a atividade de coleta de água em formigas cortadeiras Atta sexdens rubropilosa. O uso do modelo de pontes aqui proposto com fendas, poderá ser uma interessante ferramenta para exploração de transmissão de informação entre os membros de uma colônia The study of the division of tasks and participation in the maintenance of appropriate conditions the survival of social insects has received major contributions over the years. Allied to the degree of elaborate organization of many social insects and the presence of polymorphism in many groups with the formation of castes, division of labor makes it even more interesting to be investigated. This thesis presents the results of three experiments with colonies of leaf-cutting ants Atta sexdens rubropilosa: 1) it was checked if there are water marked trophallaxis between nest mates leaf-cutting ants Atta sexdens rubropilosa colonies in conditions of water stress. The results showed that the trophallaxis is an event that often occurs in colonies in water stress. 2) Was tested the division of tasks associated with the dynamics of the sizes involved in the activity of collecting water distances of 0 meter, 1 meter and 10 meters to the colonies in water stress condition. We found that after a time collecting water was associated with a variety specialized, smaller, or at the beginning of the activity of ants with larger and smaller with time ants began to perform the task, a fact that we call \"Effect of Time.\" This temporal effect of specialization varied according to the distance which the water source was placed. 3) Use the system bridges gap of 1.5 cm. A model system was developed as a bridge with gap and crevice and tested without the physical separation of sizes and the possibility of learning of the existence of the gap. The results indicate that there is a greater physical separation of the ants. Preliminary data suggest the existence of the learning gap. Thus this study sought to provide a contribution to the numerous connections that involve the activity of collecting water in l leaf-cutting ants Atta sexdens rubropilosa. The use of the bridge model proposed here with cracks may be an interesting tool for exploration of information transmission between members of a colony.

Machos dos opiliões goniosomatíneos Serracutisoma spelaeum e Serracutisoma guaricana, cujas fêmeas copulam com mais de um parceiro em um único período reprodutivo, apresentam uma grande variedade morfológica, especialmente relacionada a caracteres sexuais secundários. Considerando-se que machos menores frequentemente perdem disputas por fêmeas, esta tese objetivou verificar possíveis estratégias reprodutivas alternativas, resultantes de seleção sexual pré e pós-copulatória, adotadas por machos de pequeno porte dessas espécies assim como o papel da fêmea nesses mecanismos. Este trabalho foi, então, dividido em três capítulos, cada um com objetivos específicos. O primeiro deles refere-se à organização do trato reprodutor feminino e caracterização das espermatecas, órgãos de armazenamento de espermatozóides. Imagens de microscopia de luz e de microscopia eletrônica de varredura associadas às obtidas sob estereomicroscópio revelaram a existência de quatro espermatecas inseridas na musculatura da porção distal do ovipositor. Cada uma delas é constituída por quatro sáculos individuais de fundo cego e se comunica com a luz do ovipositor através de um único canal. A complexidade das espermatecas indica que a fêmea pode ser capaz de armazenar separadamente os espermatozóides dos diferentes machos com os quais vier a copular, enquanto a musculatura ao redor dessas estruturas sugere que ela possa determinar a paternidade de sua progênie através de contrações diferenciais. O capítulo 2 abrange observações comportamentais e características morfológicas e morfométricas do trato reprodutor masculino, objetivando identificar possíveis estratégias alternativas utilizadas por machos de pequeno porte na obtenção de sucesso reprodutivo. Para tal, foram efetuados acompanhamento do comportamento sexual e estudos morfométricos de genitália e da gônada. Machos de pequeno porte parecem adotar a estratégia de machos-satélites, copulando com as fêmeas quando machosguardiões estão com a atenção desviada. A ausência de diferenças estatísticas significativas na largura da placa ventral sugere que o pênis de machos grandes e de machos pequenos se ajustam à genitália feminina da mesma maneira e, portanto, sejam vii avaliados pelas fêmeas da mesma forma. Já a semelhança no comprimento da genitália e o menor tamanho corpóreo dos machos pequenos indicam que, durante a cópula, eles poderiam alcançar regiões mais profundas da espermateca, removendo ou deslocando os espermatozóides dos outros machos e, consequentemente, diminuindo o risco de competição espermática. A análise ultraestrutural dos espermatozóides (compostos por núcleo, citoplasma e acrossomo, além de diversas projeções na superfície) mostrou que, provavelmente, todos os machos possuam espermatozóides viáveis, visto que espermatozóides não-férteis, em geral, não possuem acrossomo. As similaridades entre machos de grande e de pequeno porte tanto na morfologia dos gametas como em seu comprimento indicam que tais características possam ter sido objeto de seleção sexual pós-copulatória. O terceiro capítulo propõe uma hipótese para a formação das projeções encontradas nos espermatozóides das duas espécies e que lhes confere uma morfologia peculiar. Para isso foi utilizada a associação de três técnicas de microscopia: microscopia eletrônica de transmissão, de varredura e confocal de varredura a laser. Os resultados obtidos evidenciam que essas estruturas são formadas ao longo do processo de espermatogênese. Uma rede de proteínas situada logo abaixo da membrana plasmática exerceria uma pressão sobre o citoplasma, que extravasaria pelos espaços existentes nessa espécie de malha. Dessa forma, as projeções seriam constituídas de citoplasma envolto por membrana plasmática. Em conclusão, os resultados obtidos na presente tese sugerem que a competição espermática e a escolha críptica da fêmea, componentes da seleção sexual póscopulatória, são pressões seletivas atuantes nessas duas espécies de opilião. Males of the goniosomatine harvestmen Serracutisoma spelaeum and Serracutisoma guaricana, which females may copulate with more than one partner during the same reproductive season, show a large morphological variation, especially considering secondary sexual features. Since smaller males frequently lose fights against larger males for females, the present thesis aimed at verifying the occurrence of alternative reproductive strategies among these smaller males as a result of pre- and postcopulatory sexual selection, as well as the role of the female in these mechanisms. This study is composed of three chapters, each one with specific objectives, as described as follows. The first chapter describes the general organization of the female reproductive system and the spermathecae, structures for the storage of sperm. Ligth and scanning electronic microscopy images revealed the presence of four spermathecae inserted in the musculature of the distal portion of the ovipositor. Each spermatheca is composed of four individual dead-end sacs and is connected to the ovipositor lumen through a single channel. This complex spermathecae structure indicates that the female is able to store sperm from different males with which she copulates in different places, and that the musculature that involves these structures may be responsible for differential contractions which would result in different paternity for the females descendants. The second chapter includes behavioral observations and morphological and morphometric analysis of the male reproductive system (genitalic and gonadal features), aiming at identifying possible alternative reproductive strategies used by small males to achieve reproductive success. Small males seem to adopt a satellite-male strategy, copulating with females when larger guardian-males are not paying attention. The absence of significant statistic differences in the width of its ventral plate suggests that the penis of both large and small males may adjust to the female genitalia in the same way and, therefore, will be similarly evaluated by the female during copulation. In turn, the smaller length associated to the same sized penis of smaller males in comparison to the larger ones may indicate that they may reach deeper regions of the female genitalia during copulation, possibly removing or displacing the sperm of other males and, in consequence, reducing the risk of sperm competition. The ultrastructure analysis of the spermatozoa (which are composed of a nucleus, cytoplasm and an acrosome, in addition to several surface projections) showed that probably males of different sizes all have viable sperm, since generally non-fertile spermatozoa lack an acrosome. Similarities in the morphology and size of the gametes of both small and large males may indicate that these features have been subjected to postcopulatory sexual selection. The third chapter suggests a new hypothesis for the development of the projections found in the surface of the spermatozoa of the two species here studied. For this study, three microscope techniques were used: transmission electronic microscopy, scanning electronic microscopy, and confocal laser scanning microscopy. The results show that these projections are formed during the spermatogenesis. A net of proteins situated just below the plasmatic membrane restrains the cytoplasm, which flows through the spaces formed in the net, forming the projections, which are externally limited by plasmatic membrane. In conclusion, the results obtained in the present study suggest that sperm competition and female cryptic choice, which are components of the postcopulatory sexual selection process, are selection pressures working on the species studied herein.

Este trabalho busca entender a diversificação craniana dos macacos do Velho Mundo (Catarrhini) integrando duas abordagens para o estudo da evolução de caracteres complexos: a genética quantitativa e a integração morfológica. A investigação tem três objetivos principais: 1) comparar a magnitude e o padrão das relações entre os caracteres cranianos entre todos os Catarrhini; 2) testar a hipótese de que deriva genética é o único agente responsável pela diversificação craniana; 3) explorar as conseqüências evolutivas da associação entre caracteres. De posse de um banco de dados bastante representativo da diversidade dos macacos do Velho Mundo (39 medidas cranianas de cerca de 6.000 crânios de mais de 130 espécies), gerei as matrizes de correlação e de variância/covariância, que resumem as relações entre os caracteres, e comparei-as entre vários grupos. Comparei-as também a expectativas derivadas de modelos teóricos de evolução por deriva genética, além de simular a ação de seleção natural sobre essas matrizes para observar o comportamento evolutivo dos diversos padrões de associação entre caracteres. De maneira geral, o padrão das relações é o mesmo entre todos os Catarrhini, mas a magnitude com que os caracteres estão associados varia bastante. Isso tem conseqüências evolutivas importantíssimas, pois grupos com baixas magnitudes tendem a responder na mesma direção em que a seleção atua (alta flexibilidade evolutiva), enquanto que altas magnitudes estão associadas, independentemente da direção da seleção, a respostas ao longo do eixo de maior variação, que no caso dos Catarrhini corresponde à variação no tamanho (baixa flexibilidade evolutiva). A diversificação inicial do grupo parece ter sido gerada por seleção natural, mas nos níveis de gênero e espécie, deriva genética é o processo predominante; a exceção são os cercopitecíneos, onde há evidência de seleção também nesses níveis. Com base nesses resultados, proponho um modelo que associa a magnitude geral da correlação entre caracteres aos possíveis caminhos evolutivos que uma população pode seguir. Apesar de este trabalho estar empiricamente restrito aos macacos do Velho Mundo, esse modelo é válido para os mamíferos como um todo e pode ser testado em outros grupos, aumentando nossa compreensão de como a associação entre caracteres afeta a evolução dos seres vivos. This is a study on the cranial diversification of the Catarrhini, a large group of primates that includes all Old World monkeys and apes, bringing together two approaches to investigate the evolution of complex characters: quantitative genetics and morphological integration. It has three main goals: 1) to compare magnitudes and patterns of inter-trait relationships in the skull among catarrhines; 2) to test the null hypothesis that genetic drift is the sole agent responsible for cranial diversification; 3) to explore the evolutionary consequences of inter-trait associations. With a large and representative cranial database of Old World monkeys and apes (39 measurements of around 6,000 skulls from more than 130 species), I generated and compared correlation and variance/covariance matrices, which summarize inter-trait relationships, among several Catarrhini groups. I compared some of those matrices to expectations derived from theoretical models of evolution through genetic drift, and simulated natural selection to observe the evolutionary behavior of each matrix. From a broad perspective, the patterns of relationships are the same among all catarrhines, but the magnitudes are quite variable. This has very important evolutionary consequences, because groups with low overall magnitudes tend to respond in the same direction of selection (high evolutionary flexibility), while higher magnitudes, regardless of the direction of selection, are associated to responses along the axis of highest variation, which in this case corresponds to size variation (low evolutionary flexibility). The initial diversification of catarrhines seems to have been generated by natural selection, but drift probably played a major role at the genus and species level; the exception are the cercopithecines, for which there is evidence for selection also in those levels. Based on these results, I propose a model that links the overall magnitude of inter-trait correlations to the possible evolutionary paths of a given population. This study is empirically restricted to Old World monkeys and apes, but the model has been proved valid to a broader sample of mammals and can be tested for other groups, contributing for our understanding of how complex characters evolve.