Explanation note: Data matrix contains: Chromosomal data matrix, Molecular data matrix and Combined data matrix. Mesoamerican and South American howlers were karyologically compared. FISH analysis using the chromosome painting probes for the 3 and 15 human chromosomes was applied to corroborate the homeology of the sexual systems. The autosomes involved in the translocation that formed the sexual systems in the Mesoamerican and South American species are different, thus suggesting an independent origin. Parsimony analysis resolved the phylogenetic relationships among howler species, demonstrating utility of the combined approach.
Villalobos et al. Ddl We present the first phylogenetic study using a combined analysis of chromosomal and molecular characters in Ceboidea to contribute to the characterization of the speciogenic processes in howler monkeys. Handbook of the Mammals of the World: Volume 3, Primates. There remains a lack of consensus regarding both the number of species within the genus, which, depending on the author, ranges from 9 to 14 species RylandsGroves, GregorinRylands and Mittermeierand the Bi yan relationships among them. Journal List Comp Cytogenet v. The analysis using only the partition of chromosomal data did not resolve the node Alouatta palliataAlouatta pigra Del rio sex with monkey, Alouatta carayaAlouatta belzebulAlouatta guariba clamitansAlouatta guariba guaribaAlouatta macconnelli Alouatta saraAlouatta seniculus arctoideasince it was Taylor cole and jesse mccartney in a polytomy Appendix 2: Figure Sc.
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Explanation note: Data matrix contains: Chromosomal data matrix, Molecular data matrix and Combined data matrix. Mesoamerican and South American howlers were karyologically compared. FISH analysis using the chromosome painting probes for the 3 and 15 human chromosomes was applied to corroborate the homeology of the sexual systems. The autosomes involved in the translocation that formed the sexual systems in the Mesoamerican and South American species are different, thus suggesting an independent origin.
Parsimony analysis resolved the phylogenetic relationships among howler species, demonstrating utility of the combined approach. A hypothesis for the origin of the multiple sex chromosome systems for the genus is proposed. Their distribution extends from southern Mexico to northern Argentina Crockett and Eisenberg , Rylands They inhabit a diverse range of environments, including tropical rain forests, flood forests, gallery forests, patches of forest and deciduous and semideciduous seasonal environments Crockett and Eisenberg , Zunino et al.
There remains a lack of consensus regarding both the number of species within the genus, which, depending on the author, ranges from 9 to 14 species Rylands , Groves , , Gregorin , Rylands and Mittermeier , and the phylogenetic relationships among them.
This shows the complexity of the taxonomy of Alouatta and highlights the importance of including a larger number of variables for a more accurate characterization of the species in the genus.
In this context, and to contribute to the description of the phylogenetic relationships in the genus, several authors have proposed that chromosomal data can also be used as phylogenetic markers, since they are inherited as mendelian characters and are conserved within species Sankoff , Dobigny et al. Following the Maximum Parsimony criterion, karyological comparisons allow the identification of chromosomal forms shared by common ancestrality.
In primates, different researchers in the last three decades have proposed chromosomal speciation as a probable evolutionary mechanism to explain the diversity observed in living species de Grouchy et al. In howler monkeys, species exhibit diploid numbers 2N ranging from 44 in Alouatta seniculus to 58 in Alouatta pigra , and in a large number of species, multiple sex chromosome systems in males originated from Y-autosome translocations have been described Table 1.
The chromosomes involved in the Y-autosome translocations in Alouatta caraya , Alouatta macconnelli , Alouatta guariba guariba Humboldt, , Alouatta guariba clamitans Cabrera, , Alouatta sara and Alouatta seniculus arctoidea Cabrera, , are homeologous to the same regions of human chromosomes 3 and 15 Consigliere et al. Cytogenetic characteristics of howler monkeys Alouatta. ND: not yet cytogenetically characterized.
However, the combination of different variables can improve the phylogenetic signal due to the possible common shared history of different datasets. At the same time, this combination can increase the support of a tree, since different characters evolve at particular rates and will support different parts of the tree Kluge , Whittaker et al. In the present contribution, howler species were karyologically compared and FISH analyses were carried out to corroborate the homeology of the sex chromosome systems among them.
Using these data and molecular data obtained from the literature, a phylogenetic analysis combining them in a single matrix was performed. Chromosome preparation : Peripheral blood samples were collected from all animals with previously heparinized disposable syringes. Metaphase spreads were treated with G-Wright banding Steinberg et al. Chromosomes were arranged according to previously described karyotypes using Photoshop CS Adobe and the species assignation of each specimen was corroborated.
Analysis of homeologies : For Alouatta caraya and Alouatta guariba clamitans , the homeologies with human chromosomes and the homeologies with the other South American howlers are well known Consigliere et al. The G-banded chromosomes of Alouatta pigra and Alouatta palliata were first compared with those of Alouatta caraya and Alouatta guariba clamitans. To compare homeologies, the G-banded metaphases obtained for Alouatta caraya , Alouatta guariba clamitans , Alouatta pigra and Alouatta palliata were also compared with those published for Alouatta guariba guariba de Oliveira et al.
FISH analysis with human chromosome painting probes 3 and 15 was used as a tool to confirm the identity of the sex chromosome systems in howlers. Homo sapiens HSA metaphases were used as a positive control of hybridization. Human X and Y chromosomes were also tested. Images were processed with Image Pro-Plus 4. Our results were compared with those previously described Consigliere et al.
Chromosomal dataset : We used data obtained from the comparisons of G-banding patterns and the analysis of chromosomal syntenic associations, both from the present study and from previous reports Consigliere et al. We considered the structural changes as characters. The pattern observed before and after their occurrence, i. The matrix see Appendix 1 was produced taking into consideration the characters proposed by Neusser et al.
These authors used an abbreviated nomenclature for ancestral Platyrrhini chromosome forms with a correspondence in the human karyotype. In the present contribution, for the character nomenclature, we refer directly to the human G band ideogram Table 2. New characters were obtained from our karyological comparisons and introduced in the chromosomal dataset. Human chromosome syntenic association considered as characters and used to construct the binary matrix of chromosomal homeologies among howler monkeys modified from Neusser et al.
Molecular dataset : The sequences available in GenBank for the same species used in the G-banding pattern and FISH comparisons were taken into to choose the molecular marker. The only molecular marker that fullfiled all the requirements was cyt b.
Cebus apella , from the Cebidae family, was taken as an outgroup species, since it is accepted that this species presents the most ancestral karyotype within Platyrrhini Clemente et al. Lagothrix lagotricha , also a member of the Atelidae family, was chosen as the second outgroup to test the monophyly of the group. All characters had the same weight, based on the premise that chromosome rearrangements occur by equal chance de Oliveira et al. The relative stability of nodes was assessed by bootstrap estimates Felsenstein based on iterations.
Each bootstrap replicate involved a heuristic parsimony search with 10 random taxon additions and tree-bisection reconnection TBR branch swapping. Karyological analysis : The cytogenetic characterization of the Alouatta specimens showed diploid numbers, sex chromosome systems and G-bandings patterns in agreement with the ones previously described for each species. On the left, human chromosomal bands with homeology for its corresponding ACA chromosome segment are indicated.
The boxes highlight the homeologies of the autosomes involved in the sex chromosome systems in these species a Comparison for ACA chromosomes 1 to 13 b Comparison for ACA chromosomes 14 to X 1. Chromosomal homeologies between Alouatta caraya and Alouatta palliata : The chromosomal rearrangements that could explain the homeologies were grouped in two categories: 1 Alouatta palliata chromosomes with no rearrangements with respect to Alouatta caraya chromosomes: 3, 5, 6, 7, 8, 9, 10, 13, 14, 16, 17, 19, 20, 21, 22, 24, 25 and X 1 ; 2 Alouatta palliata chromosomes with more than one rearrangement with respect to Alouatta caraya chromosomes: 1, 2, 4, 11, 12, 15, 18 and No homeologies were allocated for Alouatta palliata chromosome 26 and chromosome arms 4q and 2p using the level of resolution of the classical cytogenetic techniques applied.
Alouatta caraya chromosome 7 X 2 in males shares homeology with two Alouatta palliata chromosome pairs, 23 and 18, which are not the ones involved in the sex chromosome system in Alouatta palliata.
The Alouatta palliata chromosomal pair 19 X 2 in males shares homeology with chromosome 14 of Alouatta caraya. Chromosomal homeologies between Alouatta caraya and Alouatta pigra : The chromosomal rearrangements that could explain the homeologies were grouped in two categories: 1 Alouatta pigra chromosomes with no rearrangements with respect to Alouatta caraya chromosomes: 2, 5, 6, 7, 8, 10, 15, 16, 17, 20, 22, 23, 25, 28 and X 1 ; 2 Alouatta pigra chromosomes with more than one rearrangement with respect to Alouatta caraya chromosomes: 1, 3, 4, 9, 11, 13, 18, 19, 24, 26 and No homeologies were allocated for Alouatta pigra chromosomes 4p prox , 12, 14 and 21 using the level of resolution of the classical cytogenetic techniques applied.
Alouatta caraya chromosome 7 X 2 in males shares homeology with two Alouatta pigra chromosome pairs, 26 and 19, which are not the ones involved in the sex chromosome system in Alouatta pigra.
Alouatta pigra chromosome 17 X 2 in males shares homeology with chromosome 14 of Alouatta caraya which in turn has homeology with HSA7. Results show that Mesoamerican howlers share several human chromosomal syntenic associations with South American ones: HSA15qq Chromosomal homeologies between howlers, obtained from data both from this contribution and from previous reports.
Howler monkeys G-banded chromosomes with positive signal for the human chromosome painting probes analyzed. On the right, the hybridization pattern of human chromosomes 3, 21 and Alouatta palliata showed a pattern similar to that of Alouatta pigra therefore Figure 2 illustrates only the latter. The probe for the human X chromosome showed positive hybridization signal in X 1 of all the species analyzed.
The probe for the human Y chromosome did not hybridize in any of the howler species data not shown. The data obtained from the G-banding pattern and FISH homeologies, together with cyt b sequences obtained from previous reports, were used as the basis to perform a cladistic analysis.
Three data matrices were obtained: one including only chromosomal data, another including only molecular data and the last one including both types of characters chromosomal and molecular in a single matrix see Appendix 1.
Chromosomal partition : The analysis of chromosomal data resulted in 36 informative characters, 23 constant characters and 40 non-informative characters. The analysis using only the partition of chromosomal data did not resolve the node Alouatta palliata , Alouatta pigra , Alouatta caraya , Alouatta belzebul , Alouatta guariba clamitans , Alouatta guariba guariba , Alouatta macconnelli Alouatta sara , Alouatta seniculus arctoidea , since it was established in a polytomy Appendix 2: Figure Sc.
The analysis using only molecular data did not resolve the node Alouatta sara , Alouatta macconnelli , Alouatta seniculus arctoidea , Alouatta caraya , which was established as a polytomy different from that described from chromosomal data. Combined analysis : The heuristic analysis of the combined data showed a total of characters, of which were informative, non-informative and constant.
This type of analysis allowed us to solve all the nodes, resulting in a fully resolved tree. Next to the name of each species, the diploid number 2N and sex chromosome system is described. We present the first phylogenetic study using a combined analysis of chromosomal and molecular characters in Ceboidea to contribute to the characterization of the speciogenic processes in howler monkeys. The homoplasy distribution is likely to be different in each dataset because these are subject to different constraints.
Therefore, when different datasets are analyzed simultaneously, the signal common to all of them is more likely to overwhelm the homoplasy signal on the data Kluge In primates, few studies have compared and taken into account more than one type of character.
Bonvicino et al. Villalobos et al. However, these values 2N, FN, etc can be identical simply by chance and, if interpreted in a phylogenetic context, may be spurious indicators of relatedness Dobigny et al.
Our encoding strategy using the rearrangements as characters is quite similar to that used for morphological data but in cytogenetics one can retrieve information on the mutational event itself, something that is clearly not available to morphologists. Our combined phylogeny evidences the accuracy of this encoding strategy. Moreover, since Mesoamerican howlers Alouatta pigra and Alouatta palliata were poorly karyologically characterized at the time, data on these howlers are missing in all previous contributions.
This supports the basal grouping of the Alouatta pigra - Alouatta palliata Mesoamerican clade and the basal grouping of Alouatta belzebul among South American howlers. Taking into consideration the data obtained, a hypothesis can be proposed regarding the origin of the sex chromosome systems in the genus.
After this separation, both groups independently acquired the multiple sex chromosome systems currently observed through independent Y-autosome translocations. In males, two fissions, one in Yp ter and another in q prox of the autosomal pair involved Aq prox , followed by translocation of Aq prox to Yq-p prox , formed the new chromosome Y 1.
The Yp ter segment is lost and the proximal region of the fissioned autosome either is lost or, in certain howler species, could have given rise to microchromosomes e. The homologous autosomal pair involved in the translocation is the one now denominated X 2. In the case of South American howlers, the autosomal pair involved would share homeology with HSA3, whereas in the Mesoamerican species it would share homeology with HSA7.
The ancestral X chromosome is shown in white, the ancestral Y chromosome in light gray and the autosomal pair A in dark gray. Two fissions occurs, one in Yp ter and another in q prox of the autosomal pair involved Aq prox.
The translocation of Yq-p prox to the Aq formed the new Y 1 chromosome and the homolog of the autosomal pair involved in the translocation is now denominated X 2. The homolog to the autosomal chromosome in question is now identified as Y 2 c Simultaneous breaks in Y 1 q and Y 2 q and a translocation between Y 1 and Y 2 further explain the hybridization pattern observed in the sex chromosome systems of South American howlers.
A de novo centromere arises in the remains of the old Y 1 now Y 2. The remains of the old Y 2 could either be lost or remain as a microchromosome in some howlers d Hybridization pattern in South American howlers. In the case of South American howlers, the new autosomal pair involved in the sex chromosome system would share homeology with HSA This last hypothesis would require a centromeric activation in Y 2.
It can be considered that multiple sex chromosome systems would be an extremely rare phenomenon due to complication in meiosis.
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Cebus Sapajus. They are readily identified as the " organ grinder " monkey, and have been used in many movies and television shows. The range of capuchin monkeys includes Central America and South America as far south as northern Argentina.
In Central America, they usually occupy the wet lowland forests on the Caribbean coast of Costa Rica and Panama and deciduous dry forest on the Pacific coast. The word "capuchin" derives from a group of friars named the Order of Friars Minor Capuchin , an offshoot from the Franciscans , who wear brown robes with large hoods. When Portuguese explorers reached the Americas in the 15th century, they found small monkeys whose coloring resembled these friars, especially when in their robes with hoods down, and named them capuchins.
The species-level taxonomy of this genus remains highly controversial, and alternative treatments than the one listed below have been suggested. In , Jessica Lynch Alfaro et al. According to genetic studies led by Lynch Alfaro in , the gracile and robust capuchins diverged approximately 6. Lynch Alfaro suspects that the divergence was triggered by the creation of the Amazon River, which separated the monkeys in the Amazon north of the Amazon River, which evolved into the gracile capuchins, from those in the Atlantic Forest south of the river, which evolved into the robust capuchins.
Gracile capuchins have longer limbs relative to their body size than robust capuchins. Gracile capuchins have rounder skulls, whereas robust capuchins have jaws better adapted for opening hard nuts.
Robust capuchins have crests and the males have beards. Capuchins are black, brown, buff or whitish, but their exact color and pattern depends on the species involved. On average, they weigh from 3 to 9 pounds and live up to 25 years old in their natural habitats. Like most New World monkeys, capuchins are diurnal and arboreal. With the exception of a midday nap, they spend their entire day searching for food.
At night, they sleep in the trees, wedged between branches. They are undemanding regarding their habitat and can thus be found in many differing areas. The capuchin monkey feeds on a vast range of food types, and is more varied than other monkeys in the family Cebidae.
They are omnivores , and consume a variety of plant parts such as leaves, flower and fruit, seeds, pith, woody tissue, sugarcane, bulb, and exudates, as well as arthropods , molluscs , a variety of vertebrates , and even primates. Capuchin monkeys inhabit a large range of Brazil and other parts of Latin and Central America.
Capuchin monkeys often live in large groups of 10 to 35 individuals within the forest, although they can easily adapt to places colonized by humans. Usually, a single male will dominate the group, and they have primary rights to mate with the females of their group. However, the white-headed capuchin groups are led by both an alpha male and an alpha female.
These primates are territorial animals, distinctly marking a central area of their territory with urine and defending it against intruders, though outer areas may overlap. The stabilization of group dynamics is served through mutual grooming, and communication occurs between the monkeys through various calls. They remain hidden among forest vegetation for most of the day, sleeping on tree branches and descending to the ground to find drinking water.
Capuchin females often direct most of their proceptive and mating behavior towards the alpha male. However, when the female reaches the end of her proceptive period, she may sometimes mate with up to six different subordinate males in one day. Females bear young every two years following a to day gestation. The young cling to their mother's chest until they are larger, then they move to her back. Adult male capuchins rarely take part in caring for the young. Juveniles become fully mature within four years for females and eight years for males.
In captivity, individuals have reached an age of 50 years , although natural life expectancy is only 15 to 25 years. Capuchin monkeys are clever and easy to train. As a result, they are used to help people who are quadriplegics in many developed countries. They have also become popular pets and attractions for street entertainment, and are hunted for meat by local people. Since they have a high reproductive rate and can easily adapt to their living environment, loss of the forest does not negatively impact the capuchin monkey populations as much as other species, although habitat fragmentation is still a threat.
The main predator of the tufted capuchin is the harpy eagle , which has been seen bringing several capuchins back to its nest. The capuchin is considered as the most intelligent New World monkey  and is often used in laboratories.
The tufted monkey is especially noted for its long-term tool usage,  one of the few examples of primate tool use other than by apes. Upon seeing macaws eating palm nuts , cracking them open with their beaks, this monkey will select a few of the ripest fruits, nip off the tip of the fruit and drink down the juice, then seemingly discard the rest of the fruit with the nut inside.
When these discarded fruits have hardened and become slightly brittle, the capuchins will gather them up again and take them to a large flat boulder where they have previously gathered a few river stones from up to a mile away. They will then use these stones, some of them weighing as much as the monkeys, to crack open the fruit to get to the nut inside.
Young capuchins will watch this process to learn from the older, more experienced adults but it takes them 8 years to master this. In , experiments were conducted on the ability of capuchins to use money.
During the mosquito season, they crush millipedes and rub the result on their backs. This acts as a natural insect repellent. When presented with a reflection, capuchin monkeys react in a way that indicates an intermediate state between seeing the mirror as another individual and recognizing the image as self.
Most animals react to seeing their reflection as if encountering another individual they do not recognize. An experiment with capuchins shows that they react to a reflection as a strange phenomenon, but not as if seeing a strange capuchin. With scenario 1, females appeared anxious and avoided eye-contact, while males made threatening gestures.
In scenario 2, there was little reaction by either males or females. When presented with a reflection, females gazed into their own eyes and made friendly gestures, such as lip-smacking and swaying. Males made more eye contact than with strangers or familiar monkeys, but reacted with signs of confusion or distress, such as squealing, curling up on the floor, or trying to escape from the test room.
The question of whether capuchin monkeys have a theory of mind—whether they can understand what another creature may know or think—has been neither proven nor disproven conclusively.
If confronted with a knower-guesser scenario, where one trainer can be observed to know the location of food and another trainer merely guesses the location of food, capuchin monkeys can learn to rely on the knower. Easily recognized as the " organ grinder " or " greyhound jockey " monkeys, capuchins are sometimes kept as exotic pets.
Sometimes they plunder fields and crops and are seen as troublesome by nearby human populations. They are also used as service animals, sometimes being called "nature's butlers". After being socialized in a human home as infants, the monkeys undergo extensive training before being placed with a quadriplegic. Around the house, the monkeys help out by doing tasks including fetching objects, turning lights on and off, and opening drink bottles.
Crystal the Monkey is a famous monkey actress. In , the U. Non-human primates are no longer recognized as service animals under the ADA. From Wikipedia, the free encyclopedia. Capuchin Temporal range: 6. Further information: Self-awareness.
Main article: Theory of mind. Cambridge University Press. Paris, impr. An illustrated dictionary of scientific terms. S; Finotelo, L. C; Pissinatti, A.
BMC Evol. Journal of Biogeography. Archived from the original PDF on An Argument for the Use of Sapajus and Cebus ". American Journal of Primatology. Wilson, D. Baltimore: Johns Hopkins University Press. Handbook of the Mammals of the World: Volume 3, Primates.
See also: Mendes Pontes, A. Zootaxa : 1— Folia Primatol. In Marsh LK ed. Primates in fragments. New York: Kluwer. Behavioral Ecology and Sociobiology. Neotrop Primates. Amazonian Rainforest. Monkey Jungle. Retrieved SPIE Professional. Retrieved 1 January American Anthropologist. Bibcode : PLoSO Journal of Political Economy.