Potto Research Articles

ANALYSIS OF HUNTERS' RETURN IN THREE MAJOR BUSHMEAT DEPOTS, IN SOUTH-WEST NIGERIA

Submissions of game harvested by hunters per expedition to bushmeat landing depots in Southwestern Nigeria were studied for one year. Three established bushmeat landing depots were purposively selected for the study. Olomore and Kila in Ogun State and Odo Ona in Oyo State. The three were so selected because of their distribution along almost the same line of geographical placement on a line with Kila falling in between the other two on a line. Hunters’ return of harvested game after overnight hunting to the three locations on weekly basis was identified and counted for a year. Twenty-one (21) species of Vertebrates in the class Mammalia, reptiles and birds were returned to the three bushmeat depots by hunters during the study. Location-wise, Olomore had the highest return (1,457) and the least was returned to Kila (987) both in Ogun State. Species-wise, Thryonomys swinderianus (Grasscutter) had the highest population of 2,383 animals with a mean value of 794.3 ± 180.58, followed by Cephalophus maxwelli (duiker) with a mean value of 184.0 ± 32.51. The least hunters’ returns were Bitis gabonica (Black cobra) and Perodicticus potto (Bosmans potto) with a mean value of 0.33 ± 0.58 and 0.67 ± 1.15 respectively.

Detection of a Larva of Armillifer armillatus in a Potto (Perodicticus potto) from the Republic of the Congo.

We determined the complete sequence of the mitochondrial DNA (mtDNA) of a parasite discovered between the subcutaneous tissue and the peritoneum of an African nocturnal non-human primate (NHP). The parasite and host sequences were obtained by a combination of Sanger sequencing and nanopore MinION techniques. Analyses of mtDNA gene arrangements and sequences unambiguously showed that the parasite investigated was the pentastomid Armillifer armillatus, also commonly named the tongue worm. The full-length mitochondrial genome of A. armillatus, measuring 16,706 bp in length, contains 13 protein-coding genes, 2 ribosomal RNA genes, and 22 transfer RNA genes, an arrangement identical to that of previously described pentastomid mitochondrial genomes. We describe here the second full mitochondrial genome of A. armillatus to date. To identify the NHP host, maximum likelihood phylogenetic analyses of a 441-bp fragment on the 12S rDNA gene and of a 1,140-bp fragment of the mitochondrial cytochrome b strongly support clustering with the African lorisid Perodicticus potto, a species that has rarely been reported as an intermediate host of this parasite.

Anatomical reorganization within the hand and forelimb of Perodicticus potto

Historical behavioral accounts note extraordinarily high grip strength in Perodicticus potto. Anatomical analysis of the potto’s hand and forelimb reveal several specializations that may facilitate this enhanced gripping ability, including a specialized vascular bundle (the retia mirabilia) within the wrist, a hyper‐abducted pollex, and a highly reduced second digit. The myological correlates of grip strength (i.e., the contractile potential of the hand and wrist musculature) of P. potto, however, have yet to be quantitatively evaluated. The current study contrasts the myology of the hand and forearm muscles within P. potto to a series of closely‐related strepsirrhines (Nycticebus coucang, Otolemur crassicaudatus, Hapalemur griseus, and Eulemur mongoz), as well as comparing the forearm musculature to a broader (n= 41) primate sample. Despite reports of the potto’s high grip strength, our findings demonstrate relatively below‐average force capacity (relatively low physiological cross‐sectional area) within the digital flexors of this species, which suggests that the gripping ability of this species may be less extraordinary than previously hypothesized. In addition to this observation, we report a number of unusual myological features within the potto, including the largest brachioradialis (an elbow flexor) of any taxon within our sample. Even more surprisingly, we identified an independent extensor indicis within P. potto, despite the vestigial nature of the second digit, and the absence of this muscle within ~25% of other taxa within our sample ‐ all of which possessed a more substantial second digit than the potto. These data underscore the curious nature of the potto’s antebrachial musculature, but suggest that adaptations within the potto’s forelimb may be more related to maximizing endurance grip potential than towards increasing myological force.Support or Funding InformationNational Science Foundation (IOS‐15‐57125 and BCS‐14‐40599)

Extraordinary grip strength and specialized myology in the hyper-derived hand of Perodicticus potto?

Previous behavioral reports of the African lorisid, Perodicticus potto, have speculated that these animals have an extraordinary grip strength. This ability is hypothesized to be facilitated by a range of anatomical features within the forelimb, ranging from the presence of a retia mirabilia in its wrist to the hyper-abduction of its pollex. Despite numerous behavioral reports, however, this claim of extraordinary grip strength has not been empirically substantiated. This study quantifies the physiological cross-sectional area of the digital flexor muscles within P.potto. These data are compared with a broad primate sample, including several similarly sized strepsirrhines. Contrary to expectation, we found that P.potto actually has relatively below-average digital flexor PCSA. However, we identified other myological characteristics in the upper limb of P.potto that were unexpected, including the largest brachioradialis muscle (an elbow flexor) among our primate sample, and - despite P.potto having only a vestigial second digit - an independent digital extensor indicis that is absent in almost a quarter of our primate sample.

Dental Signatures for Exudativory in Living Primates, with Comparisons to Other Gouging Mammals.

Exudativory, the consumption of gums, is an obligate or a facultative dietary niche for some primates and marsupials. Exudativory has been cited as a dietary niche that may have been present in early primates, so finding a dental signature for exudativory is highly desirable. The present study combines exudativorous lorisoids (galagos and lorises) into one sample to compare to closely related, non-exudativorous lorisoids to search for a consistent dental signature of exudativory. Linear measurements were taken from the toothcomb, P2 , M3 , upper canine, and P2 from skulls of 295 adult galagids and lorisids. Also, differential distribution of enamel on the anterior teeth was qualitatively investigated as a dental signature for gouging (a behavior that facilitates some exudativory) by micro-CT scanning one specimen each from two gougers, Nycticebus coucang and Callithrix jacchus, and two non-gougers, Perodicticus potto, and Saguinus fuscicollis. Non-primate gouging mammals, the vampire bat Desmodus rotundus and the sugar glider Petaurus breviceps, were compared to non-gouging relatives. Statistical analysis revealed that exudativorous galagos and lorises had significantly (P < 0.05) reduced M3 relative to non-exudativorous galagos and lorises. While the sample sizes for assessing enamel thickness were small, preliminary results show that gouging primates and non-primate mammals have reduced lingual enamel thickness on the anterior dentition compared to non-gouging relatives. We suggest that reduction of mastication, and, therefore, M3 dimensions are a likely dental signature for exudativory in Primates. While broader samples are needed to statistically confirm, differential distribution of enamel in the anterior dentition may also be a signature of exudativory. Anat Rec, 2019. © 2018 Wiley Periodicals, Inc. Anat Rec, 303:265-281, 2020. © 2018 American Association for Anatomy.

Sources of morbidity in lorises and pottos in North American zoos: A retrospective review, 1980-2010.

Delineating patterns of morbidity can reveal management practices in need of reassessment to improve individual welfare, as well as population health and sustainability. We reviewed medical records from 38 North American zoological institutions for 276 slender lorises, slow lorises, and pottos born between January 1, 1980 and December 31, 2010. This sample included animals identified as 116 Nycticebus pygmaeus, 84 N. coucang, 48 Loris tardigradus tardigradus, 6 L.t. nordicus (now classified as L. lydekkerianus nordicus), and 22 Perodicticus potto. Taxonomy for lorises and pottos is developing, and two of these populations (N. coucang and P. potto) likely included hybrids and/or multiple species. Our focus was to examine trends based on species and age. Across all species, whole body disease events, abnormalities of bloodwork, and dental diseases were the most common sources of morbidity. Other major sources of morbidity varied by species and included trauma, respiratory disease, and ocular disease. A recent upsurge in research has informed feeding practices for slow lorises living in human care, and a similar, evidence-based approach is needed to improve diets for other species. Given the prevalence of trauma in this sample, social needs and reproductive management practices are also important areas for further investigation. Species-level health trends reveal risk factors for individual welfare that can guide husbandry practices in zoos, as well as in sanctuaries caring for the influx of lorises and pottos rescued from the growing wildlife trade.

The Distribution of Ki-67 and Doublecortin-Immunopositive Cells in the Brains of Three Strepsirrhine Primates: Galago demidoff, Perodicticus potto, and Lemur catta

This study investigated the pattern of adult neurogenesis throughout the brains of three prosimian primate species using immunohistochemical techniques for endogenous markers of this neural process. Two species, Galago demidoff and Perodicticus potto, were obtained from wild populations in the primary rainforest of central Africa, while one species, Lemur catta, was captive-bred. Two brains from each species, perfusion-fixed with 4% paraformaldehyde, were sectioned (50 µm section thickness) in sagittal and coronal planes. Using Ki-67 and doublecortin (DCX) antibodies, proliferating cells and immature neurons were identified in the two canonical neurogenic sites of mammals, the subventricular zone of the lateral ventricle (SVZ) giving rise to the rostral migratory stream (RMS), and the subgranular zone of the dentate gyrus of the hippocampus. In addition a temporal migratory stream (TMS), emerging from the temporal horn of the lateral ventricle to supply the piriform cortex and adjacent brain regions with new neurons, was also evident in the three prosimian species. While no Ki-67-immunoreactive cells were observed in the cerebellum, DCX-immunopositive cells were observed in the cerebellar cortex of all three species. These findings are discussed in a phylogenetic context.

Morphological convergence in the pubis of slow-moving primates and xenarthrans.

Determining the functional significance of pubic rami is important for reconstructing locomotor behavior of fossil species. The slow loris pelvis, characterized by long pubic rami, is unusual among primates. Long pubic rami may be related to increasing the moment arm of the abdominal musculature during ventroflexion after the termination of hindlimb suspension, which is a common component of slow arboreal quadrupedalism (AQ). Some extant xenarthran species are also slow AQ taxa, and provide an ideal group to test hypotheses of morphologically convergent adaptations to slow AQ. A model relating abdominal moment arms to pubic morphology is tested in three genera of slow-moving xenarthrans (Bradypus, Choloepus, and Cyclopes) and two species of slow loris (Nycticebus coucang and Perodicticus potto), using a comparative sample of 37 species of primates and xenarthrans. Phylogenetic analyses of variance and regression were performed on pubic dimensions (superior and inferior pubic ramus length, pubic symphysis length). As a locomotor group, slow-moving xenarthrans and lorises share superior pubic rami that are longer than all other locomotor groups; at the species level, there is some overlap among slow AQ and non-slow-AQ taxa. Inferior pubic ramus and pubic symphysis lengths also differ according to locomotor category, but multiple comparisons among locomotor groups are non-significant. These results support the hypothesis that superior pubic ramus length is functionally related to slow, suspensory locomotion by increasing the leverage of the ventral abdominal musculature, and demonstrates morphological convergence among two phylogenetically distant groups of mammals that have evolved adaptations for slow, suspensory locomotion.

The Topographic Organization of Retinal Ganglion Cell Density and Spatial Resolving Power in an Unusual Arboreal and Slow-Moving Strepsirhine Primate, the Potto (Perodicticus potto)

The potto (Perodicticus potto) is an arboreal strepsirhine found in the rainforests of central Africa. In contrast to most primates, the potto shows slow-moving locomotion over the upper surface of branches, where it forages for exudates and crawling invertebrates with its head held very close to the substrate. Here, we asked whether the retina of the potto displays topographic specializations in neuronal density that correlate with its unusual lifestyle. Using stereology and retinal wholemounts, we measured the total number and topographic distribution of retinal ganglion cells (total and presumed parasol), as well as estimating the upper limits of the spatial resolution of the potto eye. We estimated ∼210,000 retinal ganglion cells, of which ∼7% (∼14,000) comprise presumed parasol ganglion cells. The topographic distribution of both total and parasol ganglion cells reveals a concentric centroperipheral organization with a nasoventral asymmetry. Combined with the upwardly shifted orbits of the potto, this nasoventral increase in parasol ganglion cell density enhances contrast sensitivity and motion detection skywards, which potentially assists with the detection of predators in the high canopy. The central area of the potto occurs ∼2.5 mm temporal to the optic disc and contains a maximum ganglion cell density of ∼4,300 cells/mm². We found no anatomical evidence of a fovea within this region. Using maximum ganglion cell density and eye size (∼14 mm), we estimated upper limits of spatial resolving power between 4.1 and 4.4 cycles/degree. Despite their reported reliance on olfaction to detect exudates, this level of spatial resolution potentially assists pottos with foraging for small invertebrates and in the detection of predators.

Cranial vault thickness in primates: Homo erectus does not have uniquely thick vault bones

Extremely thick cranial vaults have been noted as a diagnostic characteristic of Homo erectus since the first fossil of the species was identified, but relatively little work has been done on elucidating its etiology or variation across fossils, living humans, or extant non-human primates. Cranial vault thickness (CVT) is not a monolithic trait, and the responsiveness of its layers to environmental stimuli is unknown.We obtained measurements of cranial vault thickness in fossil hominins from the literature and supplemented those data with additional measurements taken on African fossil specimens. Total CVT and the thickness of the cortical and diploë layers individually were compared to measures of CVT in extant species measured from more than 500 CT scans of human and non-human primates.Frontal and parietal CVT in fossil primates was compared to a regression of CVT on cranial capacity calculated for extant species. Even after controlling for cranial capacity, African and Asian H. erectus do not have uniquely high frontal or parietal thickness residuals, either among hominins or extant primates. Extant primates with residual CVT thickness similar to or exceeding H. erectus (depending on the sex and bone analyzed) include Nycticebus coucang, Perodicticus potto, Alouatta caraya, Lophocebus albigena, Galago alleni, Mandrillus sphinx, and Propithecus diadema. However, the especially thick vaults of extant non-human primates that overlap with H. erectus values are composed primarily of cortical bone, while H. erectus and other hominins have diploë-dominated vault bones. Thus, the combination of thick vaults comprised of a thickened diploë layer may be a reliable autapomorphy for members of the genus Homo.

Assessment of the reproductive physiology of the potto (Perodicticus potto) through fecal hormone metabolite analyses and trans-abdominal ultrasonography

Potto (Perodicticus potto) reproductive biology has been minimally studied. Noninvasive endocrinology and ultrasonography are proven tools for reproductive assessment in other primates. In this study, we used fecal hormone metabolite analysis to monitor one adult male potto and four females at different life stages. Validated testosterone (T), estrone conjugate (EC), and progesterone (P4) enzyme immunoassays (EIA) were used to assess male testicular function and female ovarian and placental activity. The male excreted mean T concentrations of 4.72 (±1.66) μg/g feces, that did not differ (P > 0.05) over time or when paired with alternate females. Baseline concentrations of EC (range: 47.93-78.81 ng/g feces) and P4 (range: 2.29-12.46 μg/g feces) differed among adult females. Follicular phases averaged 9.1 days (±3.43, n = 30 phases), whereas luteal phases averaged 19.89 days (±9.49, n = 19 phases). Gestation length (n = 2 pregnancies) was 170 days. Gestational EC and P4 concentrations were positively correlated (pregnancy A, r (132) = 0.71; pregnancy B, r (145) = 0.76) and returned to non-pregnant luteal phase levels 3-7 days post parturition. Extreme differences between pregnant and non-pregnant EC and P4 concentrations may allow for one-sample pregnancy diagnosis. Trans-abdominal ultrasonography was validated for pregnancy diagnosis with the fetus observed between 100 and 110 days post breeding. To our knowledge, this is the first use of fecal endocrinology and ultrasonography to monitor reproductive function and pregnancy in this species, and the only study in any lorisid to measure progestagens in correlation with reproductive events.

A retrospective review of mortality in lorises and pottos in North American zoos, 1980-2010

Patterns of mortality in captive animals can reveal potentially problematic care prac- tices or other risk factors that may negatively impact animal health and population sustainability. We reviewed death records (necropsy and/or histopathology reports) for 367 lorises and pottos born between 1980 and 2010 that were housed in 33 North American zoos and related facilities. Our sample included 20 Loris tardigradus nordicus, 72 L. t. tardigradus, 109 Nycticebus coucang, 133 N. pygmaeus, and 33 Perodicticus potto. In all 5 groups, neonates (animals 20% of animals showed signs of disease included the cardiovascular and hemolymphatic, endocrine and metabolic, gastrointestinal, and immunologic systems. Our results indicate that reducing neonatal mortality is a major priority for sustaining these species in captivity, and common practices for managing groups during the peri- natal period should be carefully scrutinized. Other areas in need of further investigation are envi- ronmental contributions to neoplastic and infectious disease, and dietary contributions to chronic degenerative conditions.

A Survey of Husbandry Practices for Lorisid Primates in North American Zoos and Related Facilities

Zoos and related facilities in North America currently manage five species in the primate family Lorisidae: the greater (Nycticebus coucang), Bengal (N. bengalensis) and pygmy (N. pygmaeus) slow lorises, red slender loris (Loris tardigradus), and potto (Perodicticus potto). We used an online survey to describe institutional housing and husbandry practices for these species and assess the extent to which practices are consistent with established guidelines. Our results show that most captive lorisids are housed solitarily or in pairs. Most individuals occupy a single exhibit space in a building dedicated to nocturnal animals. Facilities are commonly meeting recommendations for abiotic exhibit design and are providing animals with an enriched environment. However, pottos and slender lorises currently occupy exhibit spaces smaller than the recommended minimum, and the impact of cleaning protocols on olfactory communication should be critically evaluated. Few facilities are taking advantage of the benefits of positive reinforcement training for promoting animal welfare. Research is greatly needed on the effects of exhibit lighting on behavior, health, and reproduction; and to determine how best to manage the social needs of lorisids with naturally dispersed social structures. Although captive populations of slender lorises, pottos, and slow lorises are declining, we suggest that improved husbandry knowledge has the potential to positively influence population sustainability and to enhance future efforts to manage the growing pygmy loris population.

Strepsirhine dichotomy from a human perspective (Primates: Lorisoidea, Lemuroidea)

Using monospecific, polyclonal antisera against 69 human plasma proteins, 128 antigenic determinants from 40 cross-reacting homologues were characterized in representatives of the prosimian genera Lemur, Eulemur, Varecia (Lemuridae) and Otolemur (Galagidae). Seventeen determinants from 16 different proteins were absent in homologues of the gagalo but were shared by lemurs and several platyrrhines, cercopithecids, and hominoids. Smaller locus samples for potto(Perodicticus potto) and slow loris (Nycticebus coucang) confirmed the more distant immunological relationship of lorises than of lemurs to anthropoids. If evolutionary rates are constant and equal in lorisiform and lemuriform prosimians, this patern of character distribution indicates strepsirhine paraphyly, lorises diverging earlier (possibly some 6 times 106 years) than lemurs from anthropoid ancestors. If this is so, lorises rather than true lemurs should be elected to root the polarity of character evolution in Primates. As an alternative, galagine proteins evolve more rapidly than lemurine homologues.

Distribution of the Potto Perodicticus potto (Primates: Lorisidae) in Eastern Africa, with a Description of a New Subspecies from Mount Kenya

The potto Perodicticus potto is a small, arboreal, nocturnal primate with an extensive, but poorly-known, distribution through tropical Africa, and a debated taxonomy. This paper (1) examines the Eastern Rift Valley as a major barrier to primate distribution in eastern Africa, (2) reviews the taxonomy of P. potto, (3) describes the distribution of the eastern potto P.p. ibeanus, (4) summarizes what is known about the body size, abundance, elevation and rainfall limits of P.p. ibeanus, (5) provides evidence for the presence of P. potto east of the Eastern Rift Valley, and (6) describes, names, and discusses a new subspecies of Perodicticus for Mount Kenya (the ‘Mount Kenya potto’ Perodicticus potto stockleyi). The geographic range for P.p. ibeanus is ca. 850 000 km2 and extends from western Democratic Republic of Congo to western Kenya. P.p. ibeanus occurs where the mean annual rainfall is between 1300–2000 mm, and is not known to be present below 600 m or above 2300 m. Densities range from <2–28 individuals/km2. P.p. stockleyi is described from one specimen that is phenotypically distinct from P.p. ibeanus and that derives from a population that is probably isolated from the nearest population of P.p. ibeanus by >175 km.

Hand‐rearing a Potto Perodicticus potto at Franklin Park Zoo, Boston

In August 2001 a Potto Perodicticus potto was born in the Tropical Forest building of Franklin Park Zoo, Boston. The neonate was removed the following day for hand‐rearing because its mother, which had a history of providing poor infant care, appeared to be neglecting it. Historically, Pottos in captivity have high rates of neonatal mortality that are frequently associated with maternal neglect. This paper gives detailed information about the methods used to hand‐rear this infant successfully. Notes on infant development are presented and the health issues encountered, and medical treatments given, during the first few months are described.

Home Range Analysis of Perodicticus potto edwardsi and Sciurocheirus cameronensis

Because the spatial arrangements of nocturnal prosimians are often used to indicate their social systems, it is important to assess the reliability of methods used to analyze ranging patterns. We compared methods of home range analysis for 2 species of nocturnal prosimians: central pottos (Perodicticus potto edwardsi) and Cross River Allen’s galagos ({Sciurocheirus cameronensis}). We conducted radio-tracking studies of 10 pottos and 8 galagos from October 1999 – November 2000 in the montane rain forests of southwest Cameroon. We calculated home ranges via minimum convex polygon (MCPs) and kernel analyses. Adult potto home ranges averaged 145.2 ha (MCPs) versus only 28.4 ha via kernel analysis; the difference is statistically significant. The mean home range of galagos is 18.3 ha via MCPs and 2.19 ha via kernel analysis; the difference is statistically significant. Neither MCP nor kernel analyses revealed a sex difference in adult home ranges for pottos and galagos. Kernel analysis gave more reliable estimates of home ranges than the minimum convex polygon method used in many studies of nocturnal prosimians. Minimum convex polygon analysis tended to overestimate the range sizes and to include many areas not traversed by the animal. We compared our findings with those from an earlier study of similar species in Gabon, where little attention was given to the home range analysis, technique. Together with studies of lemur spatial systems they highlight the importance of considering the method of home range analysis when it is to be applied to understanding social systems.

The three-dimensional structure of trabecular bone in the femoral head of strepsirrhine primates

It has been hypothesized for over a hundred years that trabecular bone plays an important structural role in the musculoskeletal system of animals and that it responds dynamically to applied loads through growth. The objectives of this study are to quantify the three-dimensional structure of femoral head trabecular bone in a sample of extant strepsirrhines and to relate patterns of interspecific variation to locomotor behavioral differences. The bone volume fraction (BV/TV) and fabric anisotropy of trabecular bone in the femoral heads of Cheirogaleus major, Avahi laniger, Galago senegalensis, Galago alleni, Loris tardigradus, Otolemur crassicaudatus, and Perodicticus potto were quantified in three dimensions using serial high-resolution X-ray computed tomography scan data. A volume based method was used to quantify the structural anisotropy in three cubic samples located inside the central portion of the femoral head. Significant structural differences were found between the predominantly leaping galagines and indriids and the nonleaping lorisines and cheirogaleids. The leapers in general have relatively anisotropic trabecular bone. The galagines display a unique pattern of decreasing bone volume and increasing anisotropy moving from the superior to the inferior half of the femoral head. By contrast, the nonleaping taxa possess relatively uniform and isotropic bone throughout the femoral head. The differences in femoral head trabecular structure among these taxa seem to be related to locomotor behavioral differences, reflecting variation in the use and loading of the hip joint during normal locomotion.

Variability of metapodials in primates with rudimentary digits: Ateles geoffroyi, Colobus guereza, and Perodicticus potto.

A tenet of evolutionary theory is that, within a species, phenotypic variability is inversely related to the intensity of stabilizing selection. A corollary is that a rudimentary or vestigial structure should be highly variable. This relationship between rudimentation and variability, however, may simply be part of a continuum, as several studies have shown that variability and size of a structure are inversely related. This study tests whether the first metacarpal (MC1) in Ateles geoffroyi and Colobus guereza and the second metacarpal (MC2) in Perodicticus potto are highly variable in their lengths relative to their other metapodials. The former two species have rudimentary thumbs, and the latter species has a rudimentary index finger. Fourteen other species of primates are included in the comparison. The results show that MC1 in A. geoffroyi and C. guereza and MC2 in P. potto are the relatively shortest first and second metapodials, respectively, in this sample of primates. However, an intraspecific analysis shows that neither MC1 in A. geoffroyi and C. guereza nor MC2 in P. potto is significantly more variable than the other metapodials. Nevertheless, an interspecific analysis shows that MC1 in A. geoffroyi and C. guereza is relatively the most variable among the first metapodials (i.e., MC1 and first metatarsal) in this study. MC2 in P. potto, however, is of relatively low variability compared with the other primates. These contrasting results are interpreted in terms of the developmental and evolutionary biology of digits.

Observations of Paternal Care in Perodicticus potto at the Cincinnati Zoo and Botanical Garden

Observations of Paternal Care in Perodicticus potto at the Cincinnati Zoo and Botanical Garden