No full-text available
To read the full-text of this research,
you can request a copy directly from the authors.
Inventory and Assessment of the Pongo (Linnaeus, 1760) Skeletal Collection Housed at the Yale Peabody Museum
Abstract
Museum collections are critical resources for examination of comparative anatomy, developmental biology, and life history hypotheses. Skeletal collections provide insight into spatiotemporal, species, population, and individual variation associated with environmental, social, and epidemiological history. For endangered species such as great apes, these collections provide data nearly impossible to replicate today. In this third in a series of articles reviewing the great ape holdings of the Yale Peabody Museum, we describe the Pongo skeletal collection. Nine catalog numbers in the collection represent nine individuals: two males, five females, and two individuals of indeterminate sex. Evidence of trauma, socioecology, metabolic stressors, infectious disease, and captive management issues are described in detail. The collection exhibits taxonomic diversity as well, with two species present. Our evaluation of the Yale Peabody Museum collection provides a baseline for future research and testable hypotheses for alternate techniques, such as isotopic analyses of dental calculus and noninvasive genetic testing. Museum collections continue to provide new insights into taxonomic and individual variation and environmental context, ultimately allowing for comparisons between modern and historical environmental and behavioral variables.
ResearchGate has not been able to resolve any citations for this publication.
Among extant great apes, orangutans are considered the most sexually dimorphic in body size. However, the expression of sexual dimorphism in orangutans is more complex than simply males being larger than females. At sexual maturity, some male orangutans develop cheek pads (flanges) while other males remain unflanged even after becoming reproductively capable. Sometimes flange development is delayed in otherwise sexually mature males for a few years. In other cases, flange development is delayed for many years or decades, with some males even spending their entire lifespan as unflanged adults. Thus, unflanged males of various chronological ages can be mistakenly identified as ‘subadults’. Unflanged adult males are typically described as “female-sized”, but this may simply reflect the fact that unflanged male body size has only ever been measured in peri-pubescent individuals. In this study, we measured the skeletons of 111 wild adult orangutans (Pongo spp.) including 20 unflanged males, 45 flanged males, and 46 females, resulting in the largest skeletal sample of unflanged males yet studied. We assessed long bone lengths (as a proxy for stature) for all 111 individuals and recorded weights-at-death, femoral head diameters, bi-iliac breadths, and long bone cross-sectional areas (as proxies for mass) for 27 of these individuals, including seven flanged males, three adults confirmed-unflanged males, and three young adult likely-unflanged males. ANOVA and Kruskal-Wallis tests with Tukey and Dunn post-hoc pairwise comparisons, respectively, showed that body sizes for young adult unflanged males are similar to those of the adult females in the sample (all p ≥ 0.09 except bi-iliac breadth), whereas body sizes for adult unflanged males ranged between those of adult flanged males and adult females for several measurements (all p < 0.001). Thus, sexually mature male orangutans exhibit body sizes that range from the female end of the spectrum to the flanged male end of the spectrum. These results exemplify that the term ‘sexual dimorphism’ fails to capture the full range of variation in adult orangutan body size. By including adult unflanged males in analyses of body size and other aspects of morphology, not as aberrations but as an expected part of orangutan variation, we may begin to shift the way that we think about features typically considered dichotomous according to biological sex.
Throughout the equatorial tropics, forest conversion to agriculture often fragments crucial primate habitat. In 30 years, 80% of the alluvial lowland forests along the Kinabatangan River in Sabah, Malaysian Borneo, have been supplanted by oil palm (Elaeis guineensis) plantations. Today, only about 20% of the former orangutan (Pongo pygmaeus morio) population remains in the region. Because most of the land is now under the tenure of agribusiness companies, we used a pragmatic approach of mixed biosocial methods and citizen science engagement of oil palm growers (N = 6) as active conservation partners to study orangutan use of the privately administered landscape between protected forest fragments. We found that 22 of 25 remanent forest patches (0.5 to 242 hectares) surveyed within plantations contained food or shelter resources useful for orangutans. Of these, 20 are in regular transitory use by wider-ranging adult male orangutans, and in 9 patches, females are resident and raising offspring isolated within oil palm plantations. These findings indicate that orangutans retain a measure of normal metapopulation dynamics necessary for viability at the landscape level despite drastic habitat modification. We found that barriers to in situ conservation in these agroforest matrices were due to the following misconceptions across sectors: 1) Good farming practices require exclusion of wildlife; 2) Orangutans seen in plantations must be “rescued” by people; and 3) Translocation is an appropriate conservation strategy, and nondetrimental to orangutans. Our exploratory study exemplifies the value of biosocial methods and collaboration with industrial-scale farmers to support primate resilience in forests fragmented by agriculture.
Sexual coercion is widespread in the animal kingdom. Its direct forms, including harassment and forced copulation, have largely been investigated as an expression of (alternative) male reproductive strategies, rather than the result of a sexual conflict between the sexes. Likewise, the frequent occurrence of forced copulations in orang-utans (Pongo spp.) has been attributed to male strategies and more recently also to concealed female fecundity. So far, however, the immediate contextual variables leading to forced copulations have rarely been examined. We compared two orang-utan populations, Suaq (Pongo abelii, Sumatra) and Tuanan (Pongo pygmaeus, Borneo), both characterized by an individual-based fission–fusion lifestyle, whereas their socioecology differs. We assessed how the occurrence of female-resisted and voluntary copulations was affected by female reproductive state, male morph (unflanged or flanged), measures of male–male competition, male–female relationship and ecological factors. Besides female reproductive state and male morph, predictors of female resistance were related to male–male competition. First, female resistance was more likely towards subordinate males who were displaced from proximity to the female by another male during that association. Second, the presence of additional flanged males increased the probability of female resistance. Third, the latency to both the arrival of another male and to the end of the association after sexual interactions was shorter if there was female resistance. We conclude that sexual coercion in orang-utans is highly dependent on the vicinity of more dominant males and can only be understood in the light of sexual conflict: While males force copulations when at risk of losing access to a female and thus follow a ‘now-or-never’ strategy, female resistance follows a ‘not-you-now’ pattern, which is ultimately consistent with an infanticide avoidance strategy.
Pronounced temporal and spatial variation in the availability of food resources can produce energetic deficits in organisms. Fruit-dependent Bornean orangutans face extreme variation in fruit availability and experience negative energy and protein balance during episodes of fruit scarcity. We evaluate the possibility that orangutans of different sexes and ages catabolize muscle tissue when the availability of fruit is low. We assess variation in muscle mass by examining the relationship between urinary creatinine and specific gravity and use the residuals as a non-invasive measure of estimated lean body mass (ELBM). Despite orangutans having a suite of adaptations to buffer them from fruit scarcity and associated caloric deficits, ELBM was lower during low fruit periods in all age-sex classes. As predicted, adult male orangutans had higher ELBM than adult females and immatures. Contrary to expectation, flanged and unflanged males did not differ significantly in ELBM. These findings highlight the precarity of orangutan health in the face of rapid environmental change and add to a growing body of evidence that orangutans are characterized by unique metabolic traits shaped by their unpredictable forest environment.
DOI: 10.17014/ijog.8.1.119-130The effect of topography, climate, soil, and geology on River Sadong sediments is related to its geochemistry. Eighteen surface sediments and five core samples were collected in replicates from six sampling sites along River Sadong, Sarawak, Malaysia. The main aim of this study is to describe the geomorphology and sedimentological characteristics of the river sediments. This was achieved by the study of the processes that form the sediments and influence their physical and chemical characteristics. The sediments were analyzed for their sedimentological characteristics. The results indicated that the studied area is enriched with organic matter, and there are no sedimentary structures in its vicinity.
Nasution A, Perwitasari-Farajallah D, Utami-Atmoko SS. 2020. The distribution and density of Tapanuli orangutans (Pongo tapanuliensis) at potential corridor locations between forest fragments in Batang Toru, North Sumatra, Indonesia. Biodiversitas 21: 5382-5388. The newly discovered Tapanuli orangutan (Pongo tapanuliensis) is in danger of being lost in a few decades due to their small population number in fragmented forest of Batang Toru. The major threats faced by this critically endangered ape, including: hunting, habitat loss, and fragmentation due to large infrastructure development and other developments in the key corridor area. The establishment of corridor and buffer zone is one such effort to maintain their population and bring them from the brink of extinction. Data on the population and distribution are needed to assess and provide baseline information for investigating the habitat condition of potential corridors and buffer zones. This study aimed to analyze population and distribution of Tapanui orangutan (Pongo tapanuliensis) in buffer zone and potential corridor of Batang Toru. Our study provides an initial dataset prior to the development of the hydrodam, which is important to understand the future post-development impact. The study was carried out in disturbed (Hutaimbaru and Bulu Mario) and undisturbed forest (Sitandiang and Hopong) of Batang Toru from June 2015 until February 2016. Orangutan nest, fig (Ficus spp.), and trees bearing fruit were recorded using a line transect method. This research showed that population and distribution of orangutan were influenced by forest conditions, threat level, and fruit availability. There were fewer nests per km2 in disturbed forest than in undisturbed forest, resulting in an orangutan density estimation to be 0.24 ind/km2, 0.14 ind/ km2 (Hutaimbaru and Bulu Mario) and 0.74 ind/km2, 0.36 ind/km2 (Sitandiang and Hopong). Number of nest in transects was influenced by the availability of fig class I and class II, but not fruit tree. The APL forest of Sitandiang is key area to connect the three populations of Tapanuli orangutan: east Batang Toru, west Batang Toru, and Sibualbuali. Furthermore, orangutan’s nests were encountered both in protected and non-protected forest, thus protection of such areas, especially areas including primary forest that harbor orangutan is essential for their long term survival. The conservation measures and action Tapanuli orangutan is alarming, thus we recommend all remaining habitat should be officially and strictly protected and collaboration work with multi-stakeholder is essential to establish the corridor as well as the strategic management plan of Tapanuli orangutan.
She likes tea, sews, draws on papers and is a self-taught master of tying and untying knots. But she is not a crafty woman of the DIY set: she is Wattana, an orangutan who lives in the Jardin des Plantes Zoo in Paris. And it is in Paris where Chris Herzfeld first encounters and becomes impressed by Wattana and her exceptional abilities with knots. In Wattana: An Orangutan in Paris Herzfeld tells not only Wattana’s fascinating story, but also the story of orangutans and other primates—including bonobos, chimpanzees, and gorillas—in captivity.
Orangutans exhibit intrasex bimaturism, a trait rare among primates. Males exist in two morphs: flanged, with large bidiscoid cheek pads on their face and a laryngeal throat pouch, and unflanged, lacking secondary sexual characteristics and displaying “developmental arrest.” Flanged males in captivity are shown to have higher levels of testosterone and cortisol than unflanged males. However, research on wild orangutan remains has a greater potential to inform differences in early life stress experiences, as zoo orangutans rarely remain developmentally arrested. Here, we use associated orangutan skins and skulls to assess flanging status, canine height (n=37), and measure an early life stress indicator, linear enamel hypoplasia (LEH) defect depth, using confocal profilometry (n=7). Of the 24 defects measured, flanged male defects were deeper (range 24.7-126.6, mean 73.4) than in unflanged adult males (range 12.1-63.5, mean 31.04) (t test p=0.026). While the majority of population-level variation in defect depth is related to how rapidly canines grow in height, when assessing intrasex variation, evidence from great apes suggests that deeper defects might reflect more severe stress events during development. Flanged and unflanged males likely have similar canine development given that canine projective crown heights are similar across all males regardless of morph (p=0.47) with no overlap between males and females (p<0.001) (means: flanged 23.15, unflanged 24.55, female 14.75). Therefore, our results may indicate an adaptive benefit to arresting development is avoiding chronic stress and its associated physiological impacts. Future directions include examining differences in microanatomical growth between morphs via histologic and surface analyses.
Understanding the habitat preference and spatial distribution for the management of medium-large primates is important for conserving and enhancing biodiversity in the most isolated and remote Batang Toru landscape, North Sumatra, Indonesia. Based on the first extensive orangutan survey dataset during 2000 and 2007, we aimed to provide microhabitat preference and distribution assessment for the new species of orangutan (Pongo tapanuliensis), a poorly known and threatened primate endemic in Indonesia. To inform future conservation measures, we develop a predictive habitat suitability map and use this map to show the current threat for Tapanuli orangutan in their habitat and as the basis of proposed of the landscape boundary in Batang Toru ecosystem. In order to identify some environmental factors affecting conservation, we analyzed the microhabitat preference of Tapanuli orangutan using maximum entropy modeling (MaxEnt). The modeled orangutan distribution map covers 1.458,06 km2 (58,52% of Batang Toru’s landscape) and reveals three distinct distribution areas. The four most important environmental predictors are the distance from the cultivation area, NDVI, mean precipitation, and distance from the secondary forest edge. The distribution of the orangutan overlap with land-use categories reveals that 42,98% of the distribution lies in protected areas, but that 15,54% lies in natural forest concessions and area for other purposes (APL). Large scale land-use masterplan is needed to provide strategies and control for future development in the possibility of land uses and management are allowed in the landscape including its conservation policies. Moreover, collaborative management strategies are needed to develop a sustainable management system. We confirmed the Batang Toru landscape as the sole of Indonesia’s biodiversity hotspots and a critical area to preserve the Tapanuli orangutan.
The primate adolescent period is characterized by a series of changes in physiology, behavior, and social relationships. Orangutans have the slowest life history and the longest period of dependency of all primates. As members of a semisolitary species with high levels of sexual coercion, adolescent female orangutans face a unique combination of challenges when achieving independence from their mother. This study examined the mating behavior of adolescent female orangutans and compared it with that of adult females to assess whether mating behavior reflects distinct strategies at these different points in the life cycle. Data were collected in Gunung Palung National Park on the island of Borneo over 20 years. Mating events from adolescent (n = 19) and adult females (n = 26) were scored and compared. Adolescent female mating events had significantly higher mating scores (indicating more proceptivity) than those of adult females (β = 1.948, p = .001). Adolescent females also engaged in elaborate sociosexual interactions with different flanged males, behaviors that were never observed during mating events of adult females. These interactions involved characteristic behavior on the part of both the adolescent females and the flanged males. Given these findings and the documentation of similar accounts of adolescent female-flanged male mating from the island of Sumatra, we propose that adolescent female orangutans display distinctive behavioral repertoires throughout the genus Pongo which serves to overcome male ambivalence toward nulliparous females, establish familiarity, and evaluate coercive tendencies in flanged males. We suggest that these behavioral patterns are an integral part of female social development in a female philopatric, but highly dispersed species where consistent social support is absent after ranging independence is achieved.
The conversion of forest to agriculture continues to contribute to the loss and fragmentation of remaining orang-utan habitat. There are still few published estimates of orang-utan densities in these heavily modified agricultural areas to inform range-wide population assessments and conservation strategies. In addition, little is known about what landscape features promote orang-utan habitat use. Using indirect nest count methods, we implemented surveys and estimated population densities of the Northeast Bornean orang-utan (Pongo pygmaeus morio) across the continuous logged forest and forest remnants in a recently salvage-logged area and oil palm plantations in Sabah, Malaysian Borneo. We then assessed the influence of landscape features and forest structural metrics obtained from LiDAR data on estimates of orang-utan density. Recent salvage logging appeared to have a little short-term effect on orang-utan density (2.35 ind/km 2 ), which remained similar to recovering logged forest nearby (2.32 ind/km 2 ). Orang-utans were also present in remnant forest patches in oil palm plantations, but at significantly lower numbers (0.82 ind/km 2 ) than nearby logged forest and salvage-logged areas. Densities were strongly influenced by variation in canopy height but were not associated with other potential covariates. Our findings suggest that orang-utans currently exist, at least in the short-term, within human-modified landscapes, providing that remnant forest patches remain. We urge greater recognition of the role that these degraded habitats can have in supporting orang-utan populations, and that future range-wide analyses and conservation strategies better incorporate data from human-modified landscapes.
Despite the accumulating evidence suggesting the importance of phenotypic plasticity in diversification and adaptation, little is known about plastic variation in primate skulls. The present study evaluated the plastic variation of the mandible in Japanese macaques by comparing wild and captive specimens. The results showed that captive individuals are square-jawed with relatively longer tooth rows than wild individuals. We also found that this shape change resembles the sexual dimorphism, indicating that the mandibles of captive individuals are to some extent masculinized. By contrast, the mandible morphology was not clearly explained by ecogeographical factors. These findings suggest the possibility that perturbations in the social environment in captivity and resulting changes of androgenic hormones may have influenced the development of mandible shape. As the high plasticity of social properties is well known in wild primates, social environment may cause the inter- and intra-population diversity of skull morphology, even in the wild. The captive-wild morphological difference detected in this study, however, can also be possibly formed by other untested sources of variation (e.g. inter-population genetic variation), and therefore this hypothesis should be validated further.
Infanticide as a male reproductive tactic is widespread across mammals, and is particularly prevalent in catarrhine primates. While it has never been observed in wild orangutans, infanticide by non-sire males has been predicted to occur due to their extremely long inter-birth intervals, semi-solitary social structure, and the presence of female counter-tactics to infanticide. Here, we report on the disappearance of a healthy four-month-old infant, along with a serious foot injury suffered by the primiparous mother. No other cases of infant mortality have been observed at this site in 30 years of study. Using photographic measurements of the injury, and information on the behavior and bite size of potential predators, we evaluate the possible causes of this injury. The context, including the behavior of the female and the presence of a new male at the time of the injury, lead us to conclude that the most likely cause of the infant loss and maternal injury was male infanticide. We suggest that in orangutans, and other species where nulliparous females are not preferred mates, these females may be less successful at using paternity confusion as an infanticide avoidance tactic, thus increasing the likelihood of infanticide of their first-born infants.
We revise the type specimens of 132 nominal species of worldwide Triphoridae stored in the Natural History Museum of the United Kingdom (NHMUK), London. We provide the species name in its original combination, followed by bibliographic details of the original description, the location of the known type material, the original description (and its translation when in Latin), a diagnosis and curatorial or nomenclatural notes. We illustrated most specimens in the type series in colour and with SEM imaging and we have added the original figure whenever possible. The specimens of Triphoris alveolatus, T. granulatus, T. suturalis and T. verrucosus, all A. Adams & Reeve, 1850, T. pfeifferi Crosse & Fischer, 1865 and T. cucullatus de Folin, 1867, previously considered type material, are not considered here belonging to the type series. Adams & Reeve’s taxa should be considered nomina dubia. The name Triphora insularum is a manuscript name by H.E.J. Biggs who deposited “types” in the NHMUK but refrained from introducing the name due to the lack of apex of the studied material. We selected lectotypes for six species (T. concors Hinds, 1843, T. maxillaris Hinds, 1843, T. fuscomaculata E.A. Smith, 1904, T. shepstonensis E.A. Smith, 1906, T. eupunctata G.B. Sowerby III, 1907, and T. rufula Watson, 1886) to stabilize the nomenclature. Finally, we illustrate original specimens (although not types) of three species described by Turton, whose type material is lost.
The Natural History Museum, London (NHM), generates and holds some of the largest global data sets relating to the biological and geological diversity of the natural world. A majority of these data were, until 2015, not widely accessible, and, even when published, were typically hard to find, poorly documented and in formats that impede discovery and integration. To better serve the bespoke needs of user communities outside and within the NHM, a dedicated data portal was developed to surface these data sets and provide a sustainable platform to encourage their citation and reuse. This paper describes the technical development of the data portal, from its inception to beta launch in December 2015, its first 2 years of operation, and future plans for the project. It outlines the development principles adopted for this prototypical project, which subsequently informed new digital project management methodologies at the NHM. The process of developing the data portal acted as a driver to implement policies necessary to encourage a culture of data sharing at the NHM.
Two rare Ramphotyphlops, described in 1934 and 1993 and known only from their respective types, are rediscovered among some unidentified, no data specimens in NMW. Brief descriptions and photographs are provided for these important specimens.
Recent decades have brought countless outbreaks of infectious disease among wildlife. These events appear to be increasing in frequency and magnitude, but to objectively evaluate whether ecosystems are experiencing rising rates of disease, scientists require historical data on disease abundance. Specimens held in natural history collections represent a chronological archive of life on Earth and may, in many cases, be the only available source of data on historical disease patterns. It is possible to extract information on past disease rates by studying trace fossils (indirect fossilized evidence of an organism's presence or activity, including coprolites or feces), sequencing ancient DNA of parasites, and examining sediment samples, mummified remains, study skins (preserved animal skins prepared by taxidermy for research purposes), liquid‐preserved hosts, and hosts preserved in amber. Such use of natural history collections could expand scientific understanding of parasite responses to environmental change across deep time (that is, over the past several centuries), facilitating the development of baselines for managing contemporary wildlife disease.
The Peabody Museum of Natural History of Yale University has a long tradition of promoting and maintaining high standards of care for their collection of more than 13 million objects. Recently, an analysis of the current environmental conditions in the three primary museum buildings: 1. Peabody Museum (PM), 2. Kline Geology Laboratory (KGL), and 3. The Class of 1954 Environmental Science Center (ESC), showed that the stability of environmental conditions and efficiency in air handling systems are poorly constrained. An analysis of energy use in climate control showed that The Class of 1954 Environmental Studies Center, the most modern Yale Peabody Museum building, is the least energy efficient of the three and one of the least energy efficient museum buildings at Yale University. The Yale Peabody Museum (YPM), in collaboration with the Yale Institute for the Preservation of Cultural Heritage (IPCH), evaluated the current climate control strategy with an eye toward achieving a more practical and responsible approach, which considers the historic character of the buildings, high cost of climate control, and sensitivity to energy sustainability. This is a key mission of the university, as outlined in the Yale Sustainability Plan 2025 (https://sustainability.yale.edu/sites/default/files/sustainability_plan_2025.pdf). The main element in the transformation to a new strategy of climate management is the assessment of climate-related risks to collections. Our assessment indicates that degradation of chemically unstable polymers is the process which dominates loss of value of those collections due to climate-related risks. It is estimated that chemical degradation processes have approximately two orders of magnitude larger impact on collection value than the degradation of pyrite and pyrite containing materials and even more than pest damage. Other climate-related processes are relatively negligible in effect. These findings allow us to change the strategy of climate control, giving priority to maintaining low temperature in rooms housing collections made of organic materials and relaxing the range of allowable relative humidity parameters. The new strategy includes a shift from climate control to climate management including: dual set-point (deadband) controllers to reduce energy consumption and lower CO2 emissions, nightly shutdowns of the air handling system serving dry collections in a modern building, broader thermal criteria, and a reduction of the amount of fresh air introduced to the buildings. These will be implemented sequentially. Progress and outcomes will be the focus of a subsequent paper.
ABSTRACT
Museum collections are critical resources for examination of comparative anatomy, developmental biology, and life history hypotheses. Skeletal collections provide insight into spatiotemporal,
species, population, and individual variation associated with environmental, social, and epidemiological history. For endangered species such as great apes, these collections provide data that are
nearly impossible to replicate today. In this second in a series of articles reviewing the great ape holdings of the Peabody Museum of Natural History at Yale University, we describe the Gorilla gorilla skeletal collection, comprising 16 individuals: 13 males, 2 females, and 1 of indeterminate sex. Evidence of socioecology, interindividual violence, hunting, captive management, predation, senescence, and infectious disease are present within the skeletal material. The collection exhibits possible taxonomic diversity as well. Our evaluation of the Yale Peabody Museum collection provides a baseline for future research and testable hypotheses for alternate techniques, such as isotopic analyses of dental calculus and noninvasive genetic testing. Museum collections continue to provide new insights into taxonomic and individual variation and environmental context, and ultimately allow for comparisons between modern and historical environmental and behavioral variables.
The past two decades have seen a proliferation in bioarchaeological literature on the identification of scurvy, a disease caused by chronic vitamin C deficiency, in ancient human remains. This condition is one of the few nutritional deficiencies that can result in diagnostic osseous lesions. Scurvy is associated with low dietary diversity and its identification in human skeletal remains can provide important contextual information on subsistence strategy, resource allocation, and human‐environmental interactions in past populations. A large and robust methodological body of work on the paleopathology of scurvy exists. However, the diagnostic criteria for this disease employed by bioarchaeologists have not always been uniform. Here we draw from previous research on the skeletal manifestations of scurvy in adult and juvenile human skeletal remains and propose a weighted diagnostic system for its identification that takes into account the pathophysiology of the disease, soft tissue anatomy, and clinical research. Using a sample of individuals from the prehistoric Atacama Desert in Northern Chile, we also provide a practical example of how diagnostic value might be assigned to skeletal lesions of the disease that have not been previously described in the literature.
Conservation benefits from understanding how adaptability and threat interact to determine a taxon’s vulnerability. Recognizing how interactions with humans have shaped taxa such as the critically endangered orangutan (Pongo spp.) offers insights into this relationship. Orangutans are viewed as icons of wild nature, and most efforts to prevent their extinction have focused on protecting minimally disturbed habitat, with limited success. We synthesize fossil, archeological, genetic, and behavioral evidence to demonstrate that at least 70,000 years of human influence have shaped orangutan distribution, abundance, and ecology and will likely continue to do so in the future. Our findings indicate that orangutans are vulnerable to hunting but appear flexible in response to some other human activities. This highlights the need for a multifaceted, landscape-level approach to orangutan conservation that leverages sound policy and cooperation among government, private sector, and community stakeholders to prevent hunting, mitigate human-orangutan conflict, and preserve and reconnect remaining natural forests. Broad cooperation can be encouraged through incentives and strategies that focus on the common interests and concerns of different stakeholders. Orangutans provide an illustrative example of how acknowledging the long and pervasive influence of humans can improve strategies to preserve biodiversity in the Anthropocene.
This article examines the geographies and politics of the global trade in orangutans in the name of science and entertainment in the first half of the twentieth century, along the background of the growing international nature protection movement. In doing so, it draws attention to historical processes that led to environmental conservation in northern Sumatra. It looks at some of the causes of the wildlife trade, linking animal-human engagement in northern Sumatra in the early twentieth century to large-scale agricultural development. During this period, wildlife traders brought shipments of orangutans from northern Sumatra to Europe and elsewhere around the world. The orangutan trade and the arrival of large numbers of apes in Europe broadly influenced material practices and decisions central to the colonial project. This included the tightening of borders and surveillance in the ports and on land, as well as the creation of wildlife protection laws and policies that regulated human relationships with the environment in colonial Indonesia.
Sexually-selected infanticide by males is widespread across primates. Maternal protection is one of many infanticide avoidance strategies employed by female primates. Bornean orangutan (Pongo pygmaeus wurmbii) mothers with younger offspring are less social with males than mothers with older offspring. Additionally, the distance between a mother and offspring decreases in the presence of male conspecifics, but not female conspecifics. We hypothesized that mothers are responsible for the change in mother-offspring proximity when males are present. Using a year of behavioral data from orangutans in Gunung Palung National Park, we tested whether the Hinde Index, a ratio of the number of approaches and leaves between two individuals, was indicative of mother or offspring proximity maintenance across different social groupings. The semi-solitary social organization of orangutans allows us to observe different social groupings. We found that the mother-offspring Hinde Index was typically indicative of offspring maintenance of proximity. However, the presence of male conspecifics was associated with an increase in the Hinde Index which indicates that mothers are responsible for the decrease in mother-offspring distance when males are present. The decrease in mother-offspring distances and increase in Hinde Index when males are present indicates that mothers react to the presence of males in a protective manner. We suggest this may be an infanticide avoidance behavior by mother orangutans.
Objectives:
Developmental stress causing enamel thinning is an important topic in primate biology. Because taxa differ in growth rates and enamel thickness, the goal is to provide a new method allowing direct comparison of prevalence and salience of enamel defects among samples.
Materials and methods:
Casts of ape teeth spanning the Late Pleistocene to Late Miocene from three site areas of increasing seasonality, equator (Sumatra) to 20° (Vietnam) and 25°N latitude (China), were examined for enamel defects among paleo-orangutans (n = 571, 222, respectively) and Lufengpithecus lufengensis (n = 198). Frequency of affected teeth and number of linear enamel hypoplasia were recorded. Defect dimensions were measured with a confocal microscope. Simple prevalence is compared to weighted prevalence (%), calculated by dividing "number of LEH from specific tooth groups" by "specific tooth sample size"; this quantity divided by "tooth-specific years of imbricational enamel formation." Defect dimensions are reduced to a dimensionless index termed "enamel deficit ratio" through dividing "daily enamel deficit" by "daily secretion rate."
Results:
Weighted prevalence increases to the North, highlighting latitudinal similarities. In contrast, "enamel deficit ratio," designed to express comparative severity of developmental stress among samples, was least in the high latitude sample and differed little between paleo-orangutan samples.
Discussion:
The actual numbers generated are not as important as efficacy of the proposed methods for other taxa. Developmental stress appears least severe in the high latitude (Lufengpithecus) sample but affects a greater proportion, compared to paleo-orangutans. Regardless of findings, the proposed solutions to improve comparability of disparate samples, yield reasonable results.
Objectives
Developmental defects of tooth enamel are associated with systemic physiological stress and have been linked to seasonal environmental factors such as rainfall, temperature, and fruit availability. Here, we evaluate whether linear enamel hypoplasia and accentuated perikymata occur with any cyclicity on lower canines and then whether cycles differ between Bornean and Sumatran orangutans.
Materials and Methods
Epoxy casts of lower canines from Pongo abelii ( n = 14) and P. pygmaeus ( n = 33) were evaluated for perikymata and dental enamel defects. Individual developmental sequences (IDSs) were generated for each canine, tracking the position of each defect in the context of continuous perikymata (time). Autocorrelation, a form of time‐series statistical analysis was run for each canine to identify whether any cyclicity of defect expression was discernable.
Results
Autocorrelation revealed cycles of defect expression within canines, but no common cycle periodicities were identified between individuals of the same species or across species. P. pygmaeus averaged more linear enamel hypoplasia per year than P. abelli , but no other comparisons (number of defects, number of perikymata between defects, and autocorrelation analysis) revealed differences between the species.
Discussion
Although no common patterns of defect expression were identified within or between P. abelli and P. pygmaeus , the potential for autocorrelation analysis is promising for primatological and paleoanthropological studies of seasonal phenomena.
Conservation strategies are rarely systematically evaluated, which reduces transparency, hinders the cost-effective deployment of resources, and hides what works best in different contexts. Using data on the iconic and critically endangered orangutan (Pongo spp.), we developed a novel spatiotemporal framework for evaluating conservation investments. We show that around USD 1 billion was invested between 2000 and 2019 into orangutan conservation by governments, nongovernmental organizations, companies, and communities. Broken down by allocation to different conservation strategies, we find that habitat protection, patrolling, and public outreach had the greatest return on investment for maintaining orangutan populations. Given the variability in threats, land-use opportunity costs, and baseline remunerations in different regions, there were differential benefits per dollar invested across conservation activities and regions. We show that although challenging from a data and analysis perspective, it is possible to fully understand the relationships between conservation investments and outcomes and the external factors that influence these outcomes. Such analyses can provide improved guidance toward a more effective biodiversity conservation. Insights into the spatiotemporal interplays between the costs and benefits driving effectiveness can inform decisions about the most suitable orangutan conservation strategies for halting population declines. Although our study focuses on the three extant orangutan species of Sumatra and Borneo, our findings have broad application for evidence-based conservation science and practice worldwide.
Historical DNA (hDNA), obtained from museum and herbarium specimens, has yielded spectacular new insights into the history of organisms. This includes documenting historical genetic erosion and extinction, discovering species new to science, resolving evolutionary relationships, investigating epigenetic effects, and determining origins of infectious diseases. However, the development of best-practices in isolating, processing, and analyzing hDNA remain under-explored, due to the substantial diversity of specimen preparation types, tissue sources, archival ages, and collecting histories. Thus, for hDNA to reach its full potential, and justify the destructive sampling of the rarest specimens, more experimental work using time-series collections, and the development of improved methods to correct for data asymmetries and biases due to DNA degradation are required.
The Gunung Palung Orangutan Project has conducted research on critically endangered wild Bornean orangutans (Pongo pygmaeus wurmbii) since 1994 in Gunung Palung National Park, West Kalimantan, Indonesia. A major goal of our broad-ranging research on orangutan behavior and ecology is to understand how the unique rainforest environment of Southeast Asia, characterized by dramatic changes in fruit productivity due to unpredictable mast fruiting, impacts orangutan behavior, physiology, and health. Much of our research has been devoted to the development of non-invasive techniques and an integrated biology approach – using hormonal assays, fecal processing, nutritional analysis, genetics, and behavioral ecology – and has led to an increased understanding of the ecological and evolutionary pressures shaping orangutan adaptations. Our results show that the extended life history and very slow reproductive rate of orangutans are adaptations to their environment. Orangutans in the Gunung Palung landscape, as elsewhere across Borneo and Sumatra, also face a series of conservation challenges, including extensive habitat loss and the illegal pet trade. We highlight how our investigations of orangutan health status, ecosystem requirements, and the assessment of orangutan density using ground and drone nest surveys have been applied to conservation efforts. We describe our project's direct conservation interventions of public education and awareness campaigns, sustainable livelihood development, establishment of village-run customary forests, investigation of the illegal pet trade, and active engagement with Indonesian government organizations. These efforts, in concert with the development of local scientific and conservation capacity, provide a strong foundation for further conservation as orangutans face a challenging future.
Objectives
A fundamental assumption in biological anthropology is that living individuals will present with different growth than non‐survivors of the same population. The aim is to address the question of whether growth and development data of non‐survivors are reflective of the biological consequences of selective mortality and/or stress.
Materials and Methods
The study compares dental development and skeletal growth collected from radiographic images of contemporary samples of living and deceased individuals from the United States (birth to 20 years) and South Africa (birth to 12 years). Further evaluation of deceased individuals is used to explore differential patterns among manners of death (MOD).
Results
Results do not show any significant differences in skeletal growth or dental development between living and deceased individuals. However, in the South African deceased sample the youngest individuals exhibited substantially smaller diaphyseal lengths than the living sample, but by 2 years of age the differences were negligible. In the US sample, neither significant nor substantial differences were found in dental development or diaphyseal length according to MOD and age (>2 years of age), though some long bones in individuals <2 years of age did show significant differences. No significant differences were noted in diaphyseal length according to MOD and age in the SA sample.
Discussion
The current findings refute the idea that contemporary deceased and living individuals would present with differential growth and development patterns through all of ontogeny as well as the assumptions linking short stature, poor environments, and MOD.
Objectives
While many attempts have been made to estimate body mass in hominins from lower limb bone dimensions, the upper limb has received far less attention in this regard. Here we develop new body mass estimation equations based on humeral articular breadths in a large modern human sample and apply them to 95 Plio‐Pleistocene specimens.
Materials and Methods
Humeral head superoinferior and total distal articular mediolateral breadths were measured in a morphologically diverse sample of 611 modern human skeletons whose body masses were estimated from bi‐iliac breadth and reconstructed stature. Reduced major axis regressions were used to compute body mass estimation equations. Consistency of the resulting estimates with those derived previously using lower limb bone equations was assessed in matched Plio‐Pleistocene individuals or samples.
Results
In the modern reference sample, the new humeral body mass estimation equations exhibit only slightly lower precision compared to the previously derived lower limb bone equations. They give generally similar estimates for Pleistocene Homo , after accounting for the different shape of the humeral head articular surface in archaic Middle and Late Pleistocene Homo , except for distal humeral estimates for Late Pleistocene specimens, which average somewhat below lower limb estimates. Humeral equations give body mass estimates for australopiths that appear much too high, except for Australopithecus sediba . A chimpanzee‐based distal humeral articular formula appears to work well for larger australopith specimens.
Discussion
The new formulae provide a more secure foundation for estimating hominin body mass from humeri than previously available equations.
Despite recent rapid advances in medical knowledge that have improved survival, conventional medical science's understanding of human health and disease relies heavily on people of European descent living in contemporary urban industrialized environments. Given that modern conditions in high-income countries differ widely in terms of lifestyle and exposures compared to those experienced by billions of people and all our ancestors over several hundred thousand years, this narrow approach to the human body and health is very limiting. We argue that preventing and treating chronic diseases of aging and other mismatch diseases will require both expanding study design to sample diverse populations and contexts, and fully incorporating evolutionary perspectives. In this paper, we first assess the extent of biased representation of industrialized populations in high profile, international biomedical journals, then compare patterns of morbidity and health across world regions. We also compare demographic rates and the force of selection between subsistence and industrialized populations to reflect on the changes in how selection operates on fertility and survivorship across the lifespan. We argue that, contrary to simplistic misguided solutions like the PaleoDiet, the hypothesis of evolutionary mismatch needs critical consideration of population history, evolutionary biology and evolved reaction norms to prevent and treat diseases. We highlight the critical value of broader sampling by considering the effects of three key exposures that have radically changed over the past century in many parts of the world—pathogen burden, reproductive effort and physical activity—on autoimmune, cardiometabolic and other mismatch diseases.
Natural History Collections are a rich source of biological data. Each specimen contains data for that species’ presence for a specific location and time, providing researchers with essential biological information. Importantly, this information can be preserved and re-evaluated for hundreds of years. To maintain specimens through time, good curation protocols are essential. The Herpetology Collection (henceforth HC) at Pittsburg State University houses 1,631 specimens, representing 181 species and subspecie collected from 23 U.S. states, Mexico and Manitoba, Canada. The majority of specimens (78.6%) were collected from the four-state area (Kansas, Missouri, Arkansas and Oklahoma). Specimens collected exclusively in Kansas comprise 56.6% of the HC, and were collected in 37 counties, with major collecting effort on Crawford, Cherokee and Bourbon counties. Spatial analysis revealed many unique and unduplicated spatiotemporal records confirming the importance of the PSU herpetology collection as a local repository and source of herpetofaunal data. The temporal analysis showed continual collecting from 1961 to 1970 and from 1981 to 2002 during the months of March through June. Collecting peaks occurred by month in April, by year in 1964, and by decade during the 1960s. The curatorial work this project did on the long-neglected collections was crucial to reverse degradation, it demonstrated that specimens can be curated to 21st century standards with appropriate efforts. Out of 1,631 specimens, 147 were lost during past physical moves; out of 1,484 specimens left, 221 were rehydrated (14.8%); 757 required change of preservative fluids (51.0%); and 457 specimens (30.8%) did not require further curation other than new jars. Data limitations often were present due to somewhat incomplete descriptions of locality and habitat. Collecting biases in or nearby urban areas, along roads, and in areas of known higher biodiversity levels were identified for Kansas specimens.
Deeper or more 'severe' linear enamel hypoplasia (LEH) defects are hypothesized to reflect more severe stress during development, but it is not yet clear how depth is influenced by intrinsic enamel growth patterns. Recent work documented inter- and intraspecific differences in LEH defect depth in extant great apes, with mountain gorillas having shallower defects than other taxa, and females having deeper defects than males. Here, we assess the correspondence of inter- and intraspecific defect depth and intrinsic aspects of enamel growth: enamel extension rates, outer enamel striae of Retzius angles, and linear enamel thickness. Thin sections of great ape canines (n = 40) from Gorilla beringei beringei, Gorilla gorilla gorilla, Pan troglodytes, and Pongo spp. were analyzed. Enamel extension rates were calculated within deciles of enamel-dentine junction length. Linear enamel thickness and the angle of intersection between striae of Retzius and the outer enamel surface were measured in the imbricational enamel. Mountain gorillas have faster enamel extension rates and shallower striae angles than the other taxa examined. Mountain gorillas have thinner imbricational enamel than western lowland gorillas and orangutans, but not chimpanzees. In the combined-taxon sample, females exhibit larger striae angles and thicker imbricational enamel than males. Enamel extension rates are highly negatively correlated with striae angles and LEH defect depth. Enamel growth variation corresponds with documented inter- and intraspecific differences in LEH defect depth in great ape canines. Mountain gorillas have shallower striae angles and faster extension rates than other taxa, which might explain their shallow LEH defect morphology and the underestimation of their LEH prevalence in previous studies. These results suggest that stressors of similar magnitude and timing might produce defects of different depths in one species or sex vs. another, which has implications for interpretations of stress histories in hominins with variable enamel growth patterns.
Orangutans (Pongo spp.) are reported to have extremely slow life histories, including the longest average interbirth intervals of all mammals. Such slow life history can be viable only when unavoidable mortality is kept low. Thus, orangutans’ survivorship under natural conditions is expected to be extremely high. Previous estimates of orangutan life history were based on captive individuals living under very different circumstances or on small samples from wild populations. Here, we combine birth data from seven field sites, each with demographic data collection for at least 10 years (range 12-43 years) on wild orangutans to better document their life history. Using strict criteria for data inclusion, we calculated infant survival, interbirth intervals and female age at first reproduction, across species, subspecies and islands. We found an average closed interbirth interval of 7.6 years, as well as consistently very high pre-weaning survival for males and females. Female survival of 94% until age at first birth (at around age 15 years) was higher than reported for any other mammal species under natural conditions. Similarly, annual survival among parous females is very high, but longevity remains to be estimated. Current data suggest no major life history differences between Sumatran and Bornean orangutans. The high offspring survival is remarkable, noting that modern human populations seem to have reached the same level of survival only in the 20th century. The orangutans’ slow life history illustrates what can be achieved if a hominoid bauplan is exposed to low unavoidable mortality. Their high survival is likely due to their arboreal and non- gregarious lifestyle, and has allowed them to maintain viable populations, despite living in low- productivity habitats. However, their slow life history also implies that orangutans are highly vulnerable to a catastrophic population crash in the face of drastic habitat change.
A recent report, published by the Government of Indonesia with support from the Food and Agricultural Organization and Norway's International Climate and Forest Initiative, states that orangutan populations (Pongo spp.) have increased by more than 10% in Indonesia from 2015 to 2017, exceeding the government target of an annual 2% population increase [1]. This assessment is in strong contrast with recent publications that showed that the Bornean orangutan (P. pygmaeus) lost more than 100,000 individuals in the past 16 years [2] and declined by at least 25% over the past 10 years [3]. Furthermore, recent work has also demonstrated that both Sumatran orangutans (P. abelii) and the recently described Tapanuli orangutan (P. tapanuliensis) lost more than 60% of their key habitats between 1985 and 2007, and ongoing land use changes are expected to result in an 11–27% decline in their populations by 2020 [4,5]. Most scientific data indicate that the survival of these species continues to be seriously threatened by deforestation and killing [4,6,7] and thus all three are Critically Endangered under the International Union for Conservation of Nature's Red List. Meijaard et al. critique a 2018 report by the Indonesian government stating that orangutan populations have increased by >10%. The report contradicts recent scientific publications indicating sharp declines in all orangutan species. Collaboration between scientists and government in development and implementation of monitoring methods is advised.
Objectives: A number of studies have demonstrated the ontogenetic plasticity of long bone
diaphyseal structure in response to mechanical loading. Captivity should affect mechanical loading of the limbs, but whether captive apes grow differently than wild apes has been debated. Here, we compare captive and wild juvenile and adult Gorilla to ascertain whether growth trajectories in cross-sectional diaphyseal shape are similar in the two environments.
Materials and methods: A sample of young juvenile (n = 4) and adult (n = 10) captive Gorilla
gorilla gorilla specimens, with known life histories, were compared with age-matched wild G. g. gorilla (n = 62) and G. beringei beringei (n = 75) in relative anteroposterior to mediolateral bending strength of the femur, tibia, and humerus. Cross sections were obtained using peripheral quantitative CT.
Results: Captive and wild adult G. g. gorilla differed in bending strength ratios for all three bones, but these differences were not present in young juvenile G. g. gorilla. In comparisons across taxa, captive juvenile G. g. gorilla were more similar to wild G. g. gorilla than to G.b. beringei, while captive adult G. g. gorilla were more similar in shape to G. b. beringei in the hind limb.
Discussion: Captive and wild G. gorilla follow different ontogenetic trajectories in long bone
diaphyseal shape, corresponding to environmental differences and subsequent modified locomotor behaviors. Differences related to phylogeny are most evident early in development.
There are many reasons why museum collections may be used for destructive sampling, from DNA and isotope analysis to radiocarbon dating. The process is invasive and destroys a part, or all, of the specimen. This can result in reluctance by museum staff to allow specimens to be used in particular types of scientific research. We will present some of the motivations on both sides, but argue that the benefits of destructive sampling can outweigh the risks. Many analytical methods have improved dramatically in the last 30 years, requiring smaller sample sizes. With a focus on destructive sampling for genetic analysis, we will also present some examples from the literature where DNA from museum and archaeological specimens has greatly aided the reconstruction of a species' evolutionary history as well as enriching our understanding of the object sampled. In addition, we highlight the need for museum staff to understand exactly what researchers are asking for, and for researchers in turn to understand museum procedures. We include an example of a Destructive Sampling Policy and a Destructive Sampling Request Form, for institutions to adapt for their own use.
Nater, et al.[1] recently identified a new orangutan species (Pongo tapanuliensis) in northern Sumatra, Indonesia-just the seventh described species of living great ape. The population of this critically-endangered species is perilously small, at only ∼800 individuals [1], ranking it among the planet's rarest fauna. We assert that P. tapanuliensis is highly vulnerable to extinction because its remaining habitat is small, fragmented, and poorly protected. While road incursions within its habitat are modest-road density is only one-eighth that of northern Sumatra-over one-fifth of its habitat is zoned for agricultural conversion or is comprised of mosaic agricultural and regrowth/degraded forest. Additionally, a further 8% will be affected by flooding and infrastructure development for a hydroelectric project. We recommend urgent steps to increase the chance that P. tapanuliensis will persist in the wild.
Objective:
Linear enamel hypoplasia (LEH) is a condition marked by localized reductions in enamel thickness, resulting from growth disruptions during dental development. We use quantitative criteria to characterize the depth of LEH defects and “normal” perikymata in great apes. We test the hypothesis that mountain gorillas have shallow defects compared to other taxa, which may have led to their underestimation in previous studies.
Materials and Methods:
Previous attempts to characterize LEH morphology quantitatively have been limited in sample size and scope. We generated digital elevation models using optical profilometry (Sensofar PLu Neox) and extracted 2D coordinates using ImageJ to quantify depths in canines from three great ape genera (N = 75 perikymata; 255 defects).
Results:
All defect depths fall outside the distribution of perikymata depths. Mountain gorilla defects are significantly shallower than those of other great ape taxa examined, including western lowland gorillas. Females have significantly deeper defects than males in all taxa. The deepest defect belongs to a wild-captured zoo gorilla. Virunga mountain gorilla specimens collected by Dian Fossey exhibit deeper defects than those collected recently.
Discussion:
Shallow defect morphology in mountain gorillas may have led to an underestimation of LEH prevalence in past studies. Defect depth is used as a proxy for insult severity, but depth might be influenced by inter- and intra-specific variation in enamel growth. Future studies should test whether severe insults are associated with deeper defects, as might be the case with Haloko, a wild-captured gorilla. Ongoing histologic studies incorporating associated behavioral records will test possible factors that underlie differences in defect morphology.
Epiphyseal fusion in primates is a process that occurs in a regular sequence spanning a period of years and thus provides biological anthropologists with a useful marker of maturity that can be used to assess age and stage of development. Despite the many studies that have catalogued fusion timing and sequence pattern, comparatively little research has been devoted to understanding why these sequences exist in the first place. Answering this question is not necessarily intuitive; indeed, given that neither taxonomic affinities nor recent adaptations have been clearly defined, it is a challenge to explain this process in evolutionary terms. In all mammals, there is a tendency for the fusion of epiphyses at joints to occur close in sequence, and this has been proposed to relate to locomotor adaptations. Further consideration of the evidence suggests that linking locomotor behavior to sequence data alone is difficult to prove and may require a different type of evidence. Epiphyseal fusion should be considered in the context of other parameters that affect the developing skeleton, including how joint morphology relates to growth in length, as well as other possible morphological constraints. In recent years, developmental biology has been providing a better understanding of the molecular regulators of epiphyseal fusion. At some point in the near future, we may be able to link our understanding of the genetics of fusion timing to the possible selective mechanisms that are responsible for these sequences.
Assessing the relative contributions of immigration and diversification into the buildup of species diversity is key to understanding the role of historical processes in driving biogeographical and diversification patterns in species-rich regions. Here, we investigated how colonization, in situ speciation, and extinction history may have generated the present-day distribution and diversity of Goura crowned pigeons (Columbidae), a group of large forest-dwelling pigeons comprising four recognized species that are all endemic to New Guinea. We used a comprehensive geographical and taxonomic sampling based mostly on historical museum samples, and shallow shotgun sequencing, to generate complete mitogenomes, nuclear ribosomal clusters and independent nuclear conserved DNA elements. We used these datasets independently to reconstruct molecular phylogenies. Divergence time estimates were obtained using mitochondrial data only. All analyses revealed similar genetic divisions within the genus Goura and recovered as monophyletic groups the four species currently recognized, providing support for recent taxonomic changes based on differences in plumage characters. These four species are grouped into two pairs of strongly supported sister species, which were previously not recognized as being close relatives: Goura sclaterii with Goura cristata, and Goura victoria with Goura scheepmakeri. While the geographical origin of the Goura lineage remains elusive, the crown age of 5.73 Ma is consistent with present-day species diversity being the result of a recent diversification within New Guinea. Although the orogeny of New Guinea's central cordillera must have played a role in driving diversification in Goura, cross-barrier dispersal seems more likely than vicariance to explain the speciation events having led to the four current species. Our results also have important conservation implications. Future assessments of the conservation status of Goura species should consider threat levels following the taxonomic revision proposed by del Hoyo and Collar (HBW and BirdLife International illustrated checklist of the birds of the world 1: non-passerines, 2014), which we show to be fully supported by genomic data. In particular, by distinguishing G. sclaterii from G. scheepmakeri seems to be particularly relevant.