Limb kinematics during locomotion in the two-toed sloth (Choloepus didactylus,Xenarthra) and its implications for the evolution of the sloth locomotor apparatus

In order to gain insight into the function of the extant sloth locomotion and its evolution, we conducted a detailed videoradiographic analysis of two-toed sloth locomotion (Xenarthra: Choloepus didactylus). Both unrestrained as well as steady-state locomotion was analyzed. Spatio-temporal gait parameters, data on interlimb coordination, and limb kinematics are reported. Two-toed sloths displayed great variability in spatio-temporal gait parameters over the observed range of speeds. They increase speed by decreasing the durations of contact and swing phases, as well as by increasing step length. Gait utilization also varies with no strict gait sequence or interlimb timing evident in slow movements, but a tendency to employ diagonal sequence, diagonal couplet gaits in fast movements. In contrast, limb kinematics were highly conserved with respect to ‘normal’ pronograde locomotion. Limb element and joint angles at touch down and lift off, element and joint excursions, and contribution to body progression of individual elements are similar to those reported for non-cursorial mammals of small to medium size. Hands and feet are specialized to maintain firm connection to supports, and do not contribute to step length or progression. In so doing, the tarsometatarsus lost its role as an individual propulsive element during the evolution of suspensory locomotion. Conservative kinematic behavior of the remaining limb elements does not preclude that muscle recruitment and neuromuscular control for limb pro- and retraction are also conserved. The observed kinematic patterns of two-toed sloths improve our understanding of the convergent evolution of quadrupedal suspensory posture and locomotion in the two extant sloth lineages.     

A syndrome of mutualism reinforces the lifestyle of a sloth

Arboreal herbivory is rare among mammals. The few species with this lifestyle possess unique adaptions to overcome size-related constraints on nutritional energetics. Sloths are folivores that spend most of their time resting or eating in the forest canopy. A three-toed sloth will, however, descend its tree weekly to defecate, which is risky, energetically costly and, until now, inexplicable. We hypothesized that this behaviour sustains an ecosystem in the fur of sloths, which confers cryptic nutritional benefits to sloths. We found that the more specialized three-toed sloths harboured more phoretic moths, greater concentrations of inorganic nitrogen and higher algal biomass than the generalist two-toed sloths. Moth density was positively related to inorganic nitrogen concentration and algal biomass in the fur. We discovered that sloths consumed algae from their fur, which was highly digestible and lipid-rich. By descending a tree to defecate, sloths transport moths to their oviposition sites in sloth dung, which facilitates moth colonization of sloth fur. Moths are portals for nutrients, increasing nitrogen levels in sloth fur, which fuels algal growth. Sloths consume these algae-gardens, presumably to augment their limited diet. These linked mutualisms between moths, sloths and algae appear to aid the sloth in overcoming a highly constrained lifestyle.     

A mechanical link model of two-toed sloths: no pendular mechanics during suspensory locomotion

Sloths are morphologically specialized in suspensory quadrupedal locomotion and posture. During steady-state locomotion they utilize a trot-like footfall sequence. Contrasting the growing amount of published accounts of the functional morphology and kinematics of sloth locomotion, no study concerned with the dynamics of their quadrupedal suspensory locomotion has been conducted. Brachiating primates have been shown to travel at low mechanical costs using pendular mechanics, but this is associated with considerable dynamic forces exerted onto the support. To test whether sloth locomotion can be described by simple connected pendulum mechanics, we analyzed the dynamics of sloth locomotion with use of a mechanical segment link model. The model integrates the body segment parameters and is driven by kinematic data with both segment parameters and kinematic data obtained from the same sloth individual. No simple pendular mechanics were present. We then used the model to carry out an inverse dynamic analysis. The analysis allowed us to estimate net limb joint torques and substrate reaction forces during the contact phases. Predominant flexing limb joint torque profiles in the shoulder, elbow, hip, and knee are in stark contrast to published dominant extensor torques in the limb joints of pronograde quadrupedal mammals. This dissimilarity likely reflects the inverse orientation of the sloth towards the gravity vector. Nevertheless, scapular pivot and shoulder seem to provide the strongest torque for progression as expected based on unchanged basic kinematic pattern previously described. Our model predicts that sloths actively reduce the dynamical forces and moments that are transmitted onto the support. We conclude that these findings reflect the need to reduce the risk of breaking supports because in this case sloths would likely be unable to react quickly enough to prevent potentially lethal falls. To achieve this, sloths seem to avoid the dynamical consequences of effective pendular mechanics.     

Estimation of hearing capabilities of Early Miocene sloths (Mammalia,Xenarthra, Folivora) and palaeobiological implications

The Miocene Santacrucian sloths are a very important assemblage because they represent the first major radiation among sloths and, therefore, they could provide many insights about sloth evolution and diversity. Based on other studies, the hearing capabilities of Pleistocene sloths were identified. For a deeper understanding of these capabilities from an evolutionary point of view, in this article we study the hearing capabilities of Santacrucian sloths. We estimate the frequency range and the best frequency of hearing of six species of stem-megatherines and one species of mylodont. In most cases, the best frequency of hearing is much lower than the expected value for the estimated body mass of the species. This general shift towards low values of frequency is different than the pattern observed in mammals where sound localisation is improved by an increased sensitivity to high frequencies. In this article, we also propose some palaeobiological inferences of the low-frequency shift of hearing range in Miocene Santacrucian sloths.     

A Paleogeographic Overview of Tropical Fossil Sloths: Towards an Understanding of the Origin of Extant Suspensory Sloths?

Modern sloths are among the more characteristic mammals of South and Central American faunas. Recent discovery in four Paleogene, 22 Neogene, and dozens of Pleistocene fossiliferous localities in the tropics has revealed an unexpected paleobioversity constituted by some 81 fossil sloth species. Probably originating in southern South America near the Eocene/Oligocene transition, sloths were represented in the tropics during the late Oligocene by Pseudoglyptodon, Mylodontidae, and Megalonychidae. The latter occupied the West Indies between at least the late early Miocene and late Pleistocene, and two mylodontid clades, Octodontobradyinae and Urumacotheriinae, were characteristic of Amazonian localities from the Colhuehuapian and the Laventan periods, respectively, until the end of the Miocene. Megatheriinae and Nothrotheriidae appeared during the middle Miocene, colonizing the tropics and then North America, where Mylodontidae and Megalonychidae had already been present since the early late Miocene. Nothrotheriids are more abundant and diversified during the late Miocene in the tropics than in southern South America. Remains closely related to either of the modern sloths are absent from the fossil record, including those in the tropics. The characteristic suspensory posture of Bradypus and Choloepus appeared independently and likely after the Miocene epoch, and thus well after the hypothesized split suggested by molecular studies of the respective clades of these genera. Given their current widespread distribution in and reliance on the tropics, prospecting efforts for the direct fossil kin of suspensory sloths should concentrate on deposits in the Amazonian region, as this area has shown promise in producing fossil sloths.     

Phylogenetic relationships among sloths (Mammalia, Xenarthra, Tardigrada): the craniodental evidence

This study is undertaken in order to evaluate specific hypotheses of relationship among extant and extinct sloths (Mammalia, Xenarthra, Tardigrada). Questions of particular interest include the relationship among the three traditional family groupings of extinct ground sloths and the monophyletic or diphyletic origin of the two genera of extant tree sloths. A computer‐based cladistic investigation of the phylogenetic relationships among 33 sloth genera is performed based upon 286 osteological characteristics of the skull, lower jaw, dentition and hyoid arch. Characters are polarized via comparisons with the following successive outgroups, all members of the supraordinal grouping Edentata: the Vermilingua, or anteaters; the Cingulata, or armadillos and glyptodonts; the Palaeanodonta; and the Pholidota, or pangolins. The results of the analysis strongly corroborate the diphyly of living tree sloths, with the three‐toed sloth Bradypus positioned as the sister‐taxon to all other sloths, and the two‐toed sloth Choloepus allied with extinct members of the family Megalonychidae. These results imply that the split between the two extant sloth genera is ancient, dating back perhaps as much as 40 Myr, and that the similarities between the two taxa, including their suspensory locomotor habits, present one of the most dramatic examples of convergent evolution known among mammals. The monophyly of the three traditional ground sloth families Megatheriidae, Megalonychidae and Mylodontidae is confirmed in the present study, and the late Miocene–Pleistocene nothrotheres are shown to form a clade. It is suggested that this latter clade merits recognition as a distinct family‐level grouping, the family Nothrotheriidae. The monophyly of the Megatherioidea, a clade including members of the families Megatheriidae, Megalonychidae and Nothrotheriidae, is also supported. Within Megatherioidea, the families Nothrotheriidae and Megatheriidae form a monophyletic group called the Megatheria. The relationships within the families Megatheriidae and Mylodontidae are fully and consistently resolved, although the hypothesized scheme of relationships among the late Miocene to Pleistocene members of the mylodontid subfamily Mylodontinae differ strongly from any proposed by previous authors. Within the family Megalonychidae, Choloepus is allied to a monophyletic grouping of West Indian sloths, although the relationships within this clade are not fully resolved. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140 , 255–305.     

Effect of body orientation on regional lung expansion in dog and sloth

Recent studies (E.A. Hoffman, J. Appl. Physiol. 59: 468-480, 1985) using fast multisliced X-ray computed tomography have demonstrated a ventral-dorsal gradient of fractional lung air content (3.29% air/cm lung height) in supine dogs and an essentially uniform ventral-dorsal air content distribution in the prone dogs [mean = 66 +/- 0.6% (SE) air content]. Since the prone orientation is the dog's normal body posture, we sought to study an animal whose normal body posture was "opposite" to that of the dog. Four two-toed sloths were scanned in the Dynamic Spatial Reconstructor in the prone and supine postures. A supine fractional air content gradient was demonstrated with a regression equation of y = 2.09x + 74.3 (r = 0.92), where y is percent air content and x is vertical height in the lung, and ventral-dorsal air content distribution in the prone posture was uniform with a mean of 85 +/- 0.4% (SE) air content. The low functional residual capacity lung density in the sloth was attributable to unusually large alveoli. The mean heart volume-to-body weight ratio in the dogs was 16.4 +/- 0.6 (SE) ml/kg and that in the sloth was 7.3 +/- 0.4 (SE) ml/kg. Mean lung volume-to-body weight ratios for dogs and sloths were 57 +/- 7 (SE) and 89 +/- 6 ml/kg, respectively. Of particular interest was the fact that large changes in prone vs. supine rib cage and diaphragm geometry previously found in dogs did not occur in sloths, though significant alterations of ventral and dorsal lung geometry prone vs. supine were demonstrated, and lung shape changes in both dog and sloth are attributable to shifts in the intrathoracic position of mediastinal structures.     

Genomic evidence for rod monochromacy in sloths and armadillos suggests early subterranean history for Xenarthra

Rod monochromacy is a rare condition in vertebrates characterized by the absence of cone photoreceptor cells. The resulting phenotype is colourblindness and low acuity vision in dim-light and blindness in bright-light conditions. Early reports of xenarthrans (armadillos, sloths and anteaters) suggest that they are rod monochromats, but this has not been tested with genomic data. We searched the genomes of Dasypus novemcinctus (nine-banded armadillo), Choloepus hoffmanni (Hoffmann''s two-toed sloth) and Mylodon darwinii (extinct ground sloth) for retinal photoreceptor genes and examined them for inactivating mutations. We performed PCR and Sanger sequencing on cone phototransduction genes of 10 additional xenarthrans to test for shared inactivating mutations and estimated the timing of inactivation for photoreceptor pseudogenes. We concluded that a stem xenarthran became an long-wavelength sensitive-cone monochromat following a missense mutation at a critical residue in SWS1, and a stem cingulate (armadillos, glyptodonts and pampatheres) and stem pilosan (sloths and anteaters) independently acquired rod monochromacy early in their evolutionary history following the inactivation of LWS and PDE6C, respectively. We hypothesize that rod monochromacy in armadillos and pilosans evolved as an adaptation to a subterranean habitat in the early history of Xenarthra. The presence of rod monochromacy has major implications for understanding xenarthran behavioural ecology and evolution.     

Green sloths and brown cows: the role of dominant mammalian herbivores in carbon emissions for tropical agro‐ecosystems

When Neotropical forests are cleared, there is a rapid switch in the dominant herbivore from wild sloths (suborder Folivora) to domestic cows Bos taurus. We quantified carbon dynamics for these mammals and the ecosystems they inhabit. Because of their low metabolic rates and photosynthetically‐active algae, sloths emit trivial amounts of carbon (12 g C/sloth*day) compared to cows (2.3 kg C/cow*day). In parallel, forests are carbon sinks (?242 g C/m²*year) and pastures sources (261 g C/m²*year); cows contribute >50% of the net emissions from pastures. For a small farm in Costa Rica, this turnover in herbivores translates into ~166 metric tonnes of additional C emitted annually.     

Combined effect of ambient temperature and solar radiation on maned sloths' behaviour and detectability

Changes in ambient temperature and solar radiation may affect sloths' metabolic rate and body temperature, with consequent changes in activities, postures and microhabitat selection. Although the separate effect of temperature and solar radiation on sloth's behaviour have been previously studied, the combined effect of these climatic factors on behavioural aspects of sloths has never been systematically evaluated in field conditions. Here we evaluated the influence of hourly ambient temperature variation on maned sloth (Bradypus torquatus) activities, postures and tree crown positions, under sunny and cloudy conditions; and tested if any of the animal posture and position increase their exposure to human detection. We performed 350 h of visual observation on eight maned sloths, equipped with radio-backpacks, in northern Bahia, Brazil, recording their activities, and their resting postures and positions on tree crowns. We also recorded the time taken to visualize the sloths on 58 days to analyse if sloths' detection is affected by posture and position. Higher ambient temperature, within a range of 21–33°C, increased the sloths' activity levels in cloudy conditions but reduced their activity in sunny conditions. Increasing ambient temperature also reduced the frequency of huddled posture and increased the frequency of extended posture and permanence in the inner tree crown. Lastly, the postures and positions did not influence sloths' detectability. Thus, the direction of the temperature–activity relationship depends on climatic conditions (sunny/cloudy), and individuals rely on resting postures and positions to thermoregulate. The warmer and drier future climate, expected to occur in the northern Atlantic Forest, may impose change in the diurnal activity levels and postural pattern for this threatened species, leading maned sloths to reduce its activity on sunny and warmer days and adopting an extended posture.     

The Convergent Evolution of Suspensory Posture and Locomotion in Tree Sloths

Recent phylogenetic analyses imply a distant relationship and long separated evolution of two-toed sloths (Choloepus) and three-toed sloths (Bradypus). No known fossil sloth is interpreted to have been suspensory. As a consequence, the suspensory posture and locomotion of the extant genera likely evolved convergently in both lineages, forming a new framework for the analysis of functional aspects of the locomotor apparatus of extant tree sloths. The suspensory posture and locomotion has altered functional demands from the phylogenetically plesiomorphic non-suspensory pronograde situation. Here, anatomical traits that have been argued to be of adaptive significance for quadrupedal suspensory locomotion are reviewed and the evolution of these traits is discussed in light of the new framework. Experimental data are largely limited to Choloepus, but help to deduce functional aspects of the anatomy in Bradypus as well. The most important adaptive traits are hands and feet modified into relatively rigid hook-like appendages, great mobility of all joints proximal to the midcarpal and transverse tarsal joints, relatively long arms with a relatively short scapula, a rounded thorax with a small diameter, a highly mobile sterno-clavicular articulation, and emphasis on powerful flexion in the proximal limb joints via advantageous lever arms. Despite these changes, patterns of limb kinematics remained conservative during the course of evolution in the lineages leading to extant tree sloths, and it is suggested here that this also applies to the pattern of neuromuscular control of limb movements during locomotion. Morphological ‘solutions’ to altered functional demands posed by inversed orientation of the body differ in the two genera of extant tree sloths, thereby corroborating the proposed diphyly. Convergent evolution in tree sloths may be attributed to functional constraints posed by fossorial adaptations in early Xenarthra that canalized sloths to adopt a suspensory posture and locomotion in the arboreal habitat.     

Recent Advances on Variability, Morpho-Functional Adaptations, Dental Terminology, and Evolution of Sloths

The occasion of the Xenarthra Symposium during the ICVM 9 meeting allowed us to reflect on the considerable advances in the knowledge of sloths made by the “X-community” over the past two decades, particularly in such aspects as locomotion, mastication, diet, dental terminology, intraspecific variation, sexual dimorphism, and phylogenetic relationships. These advancements have largely been made possible by the application of cladistic methodology (including DNA analyses) and the discovery of peculiar forms such as Diabolotherium, Thalassocnus, and Pseudoglyptodon in traditionally neglected areas such as the Chilean Andes and the Peruvian Pacific desert coast. Modern tree sloths exhibit an upside-down posture and suspensory locomotion, but the habits of fossil sloths are considerably more diverse and include locomotory modes such as inferred bipedality, quadrupedality, arboreality or semiarboreality, climbing, and an aquatic or semi-aquatic lifestyle in saltwater. Modern tree sloths are generalist browsers, but fossil sloths had browsing, grazing, or mixed feeding dietary habits. Discovery of two important sloth faunas in Brazil (Jacobina) and southern North America (Daytona Beach and Rancho La Brea) have permitted evaluation of the ontogenetic variation in Eremotherium laurillardi and the existence of possible sexual dimorphism in this sloth and in Paramylodon harlani. A new dental terminology applicable to a majority of clades has been developed, facilitating comparisons among taxa. An analysis wherein functional traits were plotted onto a phylogeny of sloths was used to determine patterns of evolutionary change across the clade. These analyses suggest that megatherioid sloths were primitively semiarboreal or possessed climbing adaptations, a feature retained in some members of the family Megalonychidae. Pedolateral stance in the hindfoot is shown to be convergently acquired in Mylodontidae and Megatheria (Nothrotheriidae + Megatheriidae), this feature serving as a synapomorphy of the latter clade. Digging adaptations can only be securely ascribed to scelidotheriine and mylodontine sloths, and the latter are also the only group of grazing sloths, the remainder being general browsers.     

Three-dimensional kinematic analysis of the pectoral girdle during upside-down locomotion of two-toed sloths (Choloepus didactylus, Linné 1758)

Background

Theria (marsupials and placental mammals) are characterized by a highly mobile pectoral girdle in which the scapula has been shown to be an important propulsive element during locomotion. Shoulder function and kinematics are highly conservative during locomotion within quadrupedal therian mammals. In order to gain insight into the functional morphology and evolution of the pectoral girdle of the two-toed sloth we here analyze the anatomy and the three-dimensional (3D) pattern of shoulder kinematics during quadrupedal suspensory ('upside-down') locomotion.

Methods

We use scientific rotoscoping, a new, non-invasive, markerless approach for x-ray reconstruction of moving morphology (XROMM), to quantify in vivo the 3D movements of all constituent skeletal elements of the shoulder girdle. Additionally we use histologic staining to analyze the configuration of the sterno-clavicular articulation (SCA).

Results

Despite the inverse orientation of the body towards gravity, sloths display a 3D kinematic pattern and an orientation of the scapula relative to the thorax similar to pronograde claviculate mammalian species that differs from that of aclaviculate as well as brachiating mammals. Reduction of the relative length of the scapula alters its displacing effect on limb excursions. The configuration of the SCA maximizes mobility at this joint and demonstrates a tensile loading regime between thorax and limbs.

Conclusions

The morphological characteristics of the scapula and the SCA allow maximal mobility of the forelimb to facilitate effective locomotion within a discontinuous habitat. These evolutionary changes associated with the adoption of the suspensory posture emphasized humeral influence on forelimb motion, but allowed the retention of the plesiomorphic 3D kinematic pattern.     

Body weight: its relation to tissue composition, segment distribution, and motor function. I. Interspecific comparisons.

The composition of skin, muscle, and bone and their distribution throughout the body are compared for "advanced" or "specialized" species (Alouatta, Macaca, Canis, Felis, Lepus); smaller, more closely related species (Tupaia and the Lorisidae); and several species within the same ecosystem (Barro Colorado Island, Panama). Among the most significant variables, the skin of sloths, howlers and macaques constitutes more than 12% of body weight, whereas greyhound skin is 5% of weight; sloth and howler muscle are 25% of weight, macaque muscle about 40% of weight, greyhound and agouti muscle over 50% of weight. In tree shrews and galagos muscle is heavier (35%) than in pottos and slow lorises (below 28%), but bone and skin are lighter. All species differ in the segmental distribution of weight. Cats have light tails, light feet and heavy thighs, whereas howlers have heavy tails, heavy feet, and light thighs. The galagos have heavy hindlimbs and tails, the pottos and lorises have reduced tails and approximately equal fore- and hindlimbs. Convergences in segment pattern (sloths with pottos and lorises, marmosets with tree shrews, owl monkeys with galagos, cebus with macaques) as well as divergences are documented. All weight-of tissue and weight-of-segment variables are correlated directly with locomotor adaptation.     

Neogene sloth assemblages (Mammalia,Pilosa) of the Cocinetas Basin (La Guajira,Colombia): implications for the Great American Biotic Interchange

We describe sloth assemblages from the Cocinetas Basin (La Guajira peninsula, Colombia), found in the Neogene Castilletes and Ware formations, located in northernmost South America, documenting otherwise poorly known biotas. The tentative referral of a specimen to a small megatherioid sloth, Hyperleptus?, from the early–middle Miocene Castilletes Formation, suggests affinities of this fauna with the distant Santa Cruz Formation and documents a large latitudinal distribution for this taxon. The late Pliocene Ware Formation is much more diverse, with five distinct taxa representing every family of ‘ground sloths’. This diversity is also remarkable at the ecological level, with sloths spanning over two orders of magnitude of body mass and probably having different feeding strategies. Being only a few hundred kilometres away from the Isthmus of Panama, and a few hundred thousand years older than the classically recognized first main pulse of the Great American Biotic interchange (GABI 1), the Ware Formation furthermore documents an important fauna for the understanding of this major event in Neogene palaeobiogeography. The sloths for which unambiguous affinities were recovered are not closely related to the early immigrants found in North America before GABI 1.     

On the move: sloths and their epibionts as model mobile ecosystems

Sloths are unusual mobile ecosystems, containing a high diversity of epibionts living and growing in their fur as they climb slowly through the canopies of tropical forests. These epibionts include poorly studied algae, arthropods, fungi, and bacteria, making sloths likely reservoirs of unexplored biodiversity. This review aims to identify gaps and eliminate misconceptions in our knowledge of sloths and their epibionts, and to identify key questions to stimulate future research into the functions and roles of sloths within a broader ecological and evolutionary context. This review also seeks to position the sloth fur ecosystem as a model for addressing fundamental questions in metacommunity and movement ecology. The conceptual and evidence-based foundation of this review aims to serve as a guide for future hypothesis-driven research into sloths, their microbiota, sloth health and conservation, and the coevolution of symbioses in general.     

Molecular evidence for a diverse green algal community growing in the hair of sloths and a specific association with <Emphasis Type="Italic">Trichophilus welckeri</Emphasis>(Chlorophyta,Ulvophyceae)

Background  

Sloths are slow-moving arboreal mammals inhabiting tropical rainforests in Central and South America. The six living species of sloths are occasionally reported to display a greenish discoloration of their pelage. Trichophilus welckeri, a green algal species first described more than a century ago, is widely believed to discolor the animals fur and provide the sloth with effective camouflage. However, this phenomenon has not been explored in any detail and there is little evidence to substantiate this widely held opinion.     

Sloth Hair as a Novel Source of Fungi with Potent Anti-Parasitic,Anti-Cancer and Anti-Bacterial Bioactivity

The extraordinary biological diversity of tropical forests harbors a rich chemical diversity with enormous potential as a source of novel bioactive compounds. Of particular interest are new environments for microbial discovery. Sloths – arboreal mammals commonly found in the lowland forests of Panama – carry a wide variety of micro- and macro-organisms on their coarse outer hair. Here we report for the first time the isolation of diverse and bioactive strains of fungi from sloth hair, and their taxonomic placement. Eighty-four isolates of fungi were obtained in culture from the surface of hair that was collected from living three-toed sloths (Bradypus variegatus, Bradypodidae) in Soberanía National Park, Republic of Panama. Phylogenetic analyses revealed a diverse group of Ascomycota belonging to 28 distinct operational taxonomic units (OTUs), several of which are divergent from previously known taxa. Seventy-four isolates were cultivated in liquid broth and crude extracts were tested for bioactivity in vitro. We found a broad range of activities against strains of the parasites that cause malaria (Plasmodium falciparum) and Chagas disease (Trypanosoma cruzi), and against the human breast cancer cell line MCF-7. Fifty fungal extracts were tested for antibacterial activity in a new antibiotic profile screen called BioMAP; of these, 20 were active against at least one bacterial strain, and one had an unusual pattern of bioactivity against Gram-negative bacteria that suggests a potentially new mode of action. Together our results reveal the importance of exploring novel environments for bioactive fungi, and demonstrate for the first time the taxonomic composition and bioactivity of fungi from sloth hair.     

Unexpected Strong Polygyny in the Brown-Throated Three-Toed Sloth

Promiscuous mating strategies are much more common than previously appreciated. So much so, that several authors have proposed that promiscuity is the “rule” rather than the exception in vertebrate mating systems. Decreasing species mobility and increasing habitat fragmentation have both been suggested to reduce the “polygyny potential” of the environment and promote other mating strategies like promiscuity in females. We explored the social and genetic mating system for one of the most sedentary extant mammals, the brown-throated three-toed sloth (Bradypus variegatus), within a highly fragmented Neotropical habitat. Surprisingly, we found that three-toed sloths were strongly polygynous, with males excluding male competitors from their core ranges, and exhibiting strong reproductive skew. Indeed, only 25% of all resident adult males sired offspring and one individual sired half of all sampled juveniles. Paradoxically, a sedentary life-history strategy seems to facilitate polygyny in fragmented landscapes because multiple females can persist within small patches of habitat, and be monopolized by a single male. Our work demonstrates that strong polygyny can arise in systems in which the polygyny potential should be extremely low, and other strategies, including promiscuity, would be favoured. Mating systems can be influenced by a multitude of factor and are dynamic, varying among taxa, over time, and across habitats; consequently, mating systems remain difficult to predict based on general ecological principles.     

Multi‐scale effects of habitat structure and landscape context on a vertebrate with limited dispersal ability (the brown‐throated sloth,Bradypus variegatus)

As human population, food consumption, and demand for forest products continue to rise over the next century, the pressures of land‐use change on biodiversity are projected to intensify. In tropical regions, countryside habitats that retain abundant tree cover and structurally complex canopies may complement protected areas by providing suitable habitats and landscape connectivity for a significant portion of the native biota. Species with low dispersal capabilities are among the most at risk of extinction as a consequence of land‐use change. We assessed how the spatial distribution of the brown‐throated sloth (Bradypus variegatus), a model species for a vertebrate with limited dispersal ability, is shaped by differences in habitat structure and landscape patterns of countryside habitats in north‐central Costa Rica using a multi‐scale framework. We quantified the influence of local habitat characteristics and landscape context on sloth occurrence using mixed‐effects logistic regression models. We recorded 27 sloths within countryside habitats and found that both local and landscape factors significantly influenced their spatial distribution. Locally, sloths favored structurally complex habitats, with greater canopy cover and variation in tree height and basal area. At the landscape scale, sloths demonstrated a preference for habitats with high proportions of forest and nearby large tracts of forest. Although mixed‐use areas and tree plantations are not substitutes for protected forests, our results suggest they provide important supplemental habitats for sloths. To promote the conservation and long‐term viability of sloth populations in the tropical countryside, we recommend that land managers retain structurally complex vegetation and large patches of native habitat.