Colobus Research Articles

When population growth intensifies intergroup competition, female colobus monkeys free-ride less

Intergroup aggression often results in the production of public goods, such as a safe and stable social environment and a home range containing the resources required to survive and reproduce. We investigate temporal variation in intergroup aggression in a growing population of colobus monkeys (Colobus vellerosus) to ask a novel question: “Who stepped-up to produce these public goods when doing so became more difficult?”. Both whole-group encounters and male incursions occurred more frequently as the population grew. Males and females were both more likely to participate in whole-group encounters when monopolizable food resources were available, indicating both sexes engaged in food defence. However, only females increasingly did so as the population grew, suggesting that it was females who increasingly produced the public good of home range defence as intergroup competition intensified. Females were also more active in male incursions at high population densities, suggesting they increasingly produced the public good of a safe and stable social environment. This is not to say that males were chronic free-riders when it came to maintaining public goods. Males consistently participated in the majority of intergroup interactions throughout the study period, indicating they may have lacked the capacity to invest more time and effort.

A retrospective study on the immobilisation of five captive primate species at Uganda Wildlife Conservation Education Centre, Uganda

AbstractThis retrospective study examined data from 108 primates consisting of 41 olive baboons (Papio anubis), 30 colobus monkeys (Colobus guereza), six debrazza monkeys (Cercopithecus neglectus), 26 red‐tailed monkeys (Cercopithecus ascanius), and five ruwenzori colobus monkeys (Colobus angolensis ruwenzorii). These primates underwent annual health assessments, involving the administration of an initial intramuscular dose ketamine (5 mg/kg) plus medetomidine (0.05 mg/kg). Additional intramuscular top‐ups of ketamine (5 mg/kg) plus medetomidine (0.05 mg/kg) were given as needed. Monitoring started when the animals were placed on the examination table in dorsal recumbency and continued until their return to the enclosures. During this period, various physiological parameters, such as heart rate (beats per minute), respiration rate (breaths per minute), peripheral oxygen saturation (SpO2, %) and rectal temperature (°C), were recorded at random intervals. To reverse immobilisation, atipamezole was administered intramuscularly at five times the medetomidine dose. Additionally, the study measured various induction and recovery times.

Pump and sway: Wild primates use compliant supports as a tool to augment leaping in the canopy.

Despite qualitative observations of wild primates pumping branches before leaping across gaps in the canopy, most studies have suggested that support compliance increases the energetic cost of arboreal leaping, thus limiting leaping performance. In this study, we quantified branch pumping behavior and tree swaying in wild primates to test the hypothesis that these behaviors improve leaping performance. We recorded wild colobine monkeys crossing gaps in the canopy and quantitatively tracked the kinematics of both the monkey and the compliant support during behavioral sequences. We also empirically measured the compliance of a sample of locomotor supports in the monkeys' natural habitat, allowing us to quantify the resonant properties of substrates used during leaping. Analyses of three recordings show that adult red colobus monkeys (Piliocolobus tephrosceles) use branch compliance to their advantage by actively pumping branches before leaping, augmenting their vertical velocity at take-off. Quantitative modeling of branch resonance periods, based on empirical measurements of support compliance, suggests that monkeys specifically employed branch pumping on relatively thin branches with protracted periods of oscillation. Finally, an additional four recordings show that both red colobus and black and white colobus monkeys (Colobus guereza) utilize tree swaying to cross large gaps, augmenting horizontal velocity at take-off. This deliberate branch manipulation to produce a mechanical effect for stronger propulsion is consistent with the framework of instrumental problem-solving. To our knowledge, this is the first study of wild primates which quantitatively shows how compliant branches can be used advantageously to augment locomotor performance.

Black-and-white pelage as visually protective coloration in colobus monkeys

Conspicuous coloration is often seen as the evolutionary consequence of either sexual selection or warning predators visually about prey defences, although not all conspicuous species fit this paradigm. Exceptions include several species of colobus monkeys whose black-and-white coloration, characteristic of larger colobines, has never been explained. Here, using photographs of black-and-white colobus (Colobus vellerosus) taken in the wild, quantitative image analysis, and vision modelling (acuity-corrected feline, chimpanzee, and raptor vision), we provide evidence that their coloration provides distance-dependent camouflage in natural forest environments. At all viewing distances, black-and-white colobus monkeys blend into their environment because of their high chromatic overlap with the background when viewed by ecologically relevant predator vision models which have low visual acuities. Additionally, for chimpanzee and felid vision, there was evidence of edge disruption at longer viewing distances. Our comparative analyses of different species of colobines do not support an association between black-and-white coloration and larger body mass or group size, but this may simply be due to the limited number of species within the family. We reason that black-and-white colobines gain visually protective coloration through background matching against felids, and also benefit through disruptive coloration against felids and chimpanzees but rely on these protective coloration mechanisms less against raptors.Significance statementConspicuous black-and-white coloration in colobus monkeys has long eluded an evolutionary explanation. Our research, employing photographs of colobus monkeys, image analysis, and vision modelling, unveils a novel approach to this coloration. We show that in their natural forest habitats, black-and-white colobus monkeys employ distance-dependent camouflage against ecologically relevant predators with low visual acuity, blending in with the background at increasing viewing distances. In summary, the evolution of black-and-white colobus coloration appears to be related to background matching against felids, with added benefits of disruptive camouflage against felids and chimpanzees, less so to raptors, offering fresh insights into the complex interplay of coloration and survival strategies in mammals.

Scientific activism to protect the world's primates and their environments from extinction: Introduction to the special issue.

Nonhuman primates and their habitats are facing an impending extinction crisis. Approximately 69% of primate species are listed by the International Union for Conservation of Nature as threatened and 93% have declining populations. Human population growth (expected to reach 10.9 billion by the year 2100), the unsustainable demands of a small number of consumer nations for forest-risk commodities, deforestation and habitat conversion, the expansion of roads and rail networks, cattle ranching, the hunting and trapping of wild primate populations, and the potential spread of infectious diseases are among the primary drivers of primate population decline. Climate change will only exacerbate the current situation. The time to act to protect primate populations is now! In this special issue of the American Journal of Primatology, we present a series of commentaries, formulated as "Action Letters."These are designed to educate and inform primatologists, conservation biologists, wildlife ecologists, political leaders, and global citizens about the conservation challenges faced by particular primate taxa and particular world regions, and present examples of specific actions that one can take, individually and collectively, to promote the persistence of wild primate populations and environmental justice for local human populations and impacted ecological communities. As scientists, researchers, and educators, primatologists are in a unique position to lead local, national, and international efforts to protect biodiversity. In this special issue, we focus on primates of the Brazilian Amazon, lemurs of northeast Madagascar, Temminck's red colobus monkey (Piliocolobus badius temminckii), night monkeys (Aotus spp.), long-tailed macaques (Macaca fascicularis), the primate pet trade, and professional capacity building to foster conservation awareness and action. We encourage primatologists, regardless of their research focus, to engage in both advocacy and activism to protect wild primate populations worldwide.

Relative abundance and distribution of black-and-white colobus monkey (Colobus guereza guereza) in northwestern Ethiopia

Relative abundance and distribution of black-and-white colobus monkey (Colobus guereza guereza) in northwestern Ethiopia

Serial ultrasonographic measurements of fetal parameters over three successive pregnancies in a captive Eastern black-and-white colobus monkey (Colobus guereza).

This study provides ultrasonographic fetal growth charts for the Eastern black-and-white colobus monkey (Colobus guereza). Throughout three consecutive gestations (-162 to -2 days to parturition) in a single dam, we opportunistically obtained ultrasonographic measurements for the following parameters: biparietal diameter, head circumference, humerus length, femur length, tibia length, radius length, thoracic width, kidney length, and crown-rump length. Biparietal diameter was the most consistently measured parameter. First detection of fetuses occurred between 96 and 162 days before parturition. This report demonstrates that voluntary transabdominal ultrasound can be well-tolerated in the colobus monkey using operant conditioning. These findings may be useful to assess fetal development and predict parturition dates in the absence of a known conception date in this species.

Spinal epidural empyema in a colobus monkey

AbstractA 10‐year‐old, male colobus monkey presented with a kyphotic appearance. Lumbar spinal pain was suspected, and the colobus was started on a course of meloxicam. With no improvement noted, it was sedated for examination. Haematology revealed a neutrophilia. An empirical course of clavulanate potentiated amoxicillin was started. With no improvements seen after a 10‐day course, clinical examination was repeated. Abdominal ultrasound and abdominal radiographs revealed an abdominal effusion near the lumbar vertebrae, and haematology revealed a neutrophilia with toxic neutrophils. An enrofloxacin course was started. Three days later, the colobus presented with acute paraplegia with neurolocalisation to the T3–L3 spinal cord segments. A magnetic resonance imaging of the vertebral column was performed. Magnetic resonance imaging revealed multifocal osteolytic vertebral body lesions, compression fractures of multiple vertebral bodies and ventral spinal cord compression. Differential diagnosis included infectious osteomyelitis with spinal epidural empyema or less likely, neoplasia. Due to the poor prognosis, euthanasia was elected.

On the move: Activity budget and ranging ecology of endangered Ashy red colobus monkeys (Piliocolobus tephrosceles) in a savanna woodland habitat

On the move: Activity budget and ranging ecology of endangered Ashy red colobus monkeys (Piliocolobus tephrosceles) in a savanna woodland habitat

Wild origins and mitochondrial genetic diversity of Angolan Colobus monkeys (Colobus angolensis) in AZA-accredited zoos and its implications for ex situ population management.

Across zoo's accredited by the Association of Zoos and Aquariums(AZA), species are typically managed as a single population to retain 90% of the founding members' gene diversity. Often, little is known about the specific geographic origins of the founders or how representative the ex situ population's genetic diversity is of the wild population. This study uses mitochondrial DNA (mtDNA) sequencing to investigate haplotype diversity and geographic female founder origin of the AZA-managed Angolan colobus (Colobus angolensis) monkey population. We obtained fecal samples from individuals closely related to founder animals at five zoos and found four haplotypes among 23 individuals. Analyzed together with wild C. angolensis haplotypes, we found two haplotypes identical to those found in Tanzanian populations: one haplotype, possessed by 13 individuals (descended from three founders), matched an East Usambara Mountains haplotype, while the other, possessed by seven individuals (from four founders), matched a haplotype found in both the South Pare Mountains and Rufiji River. Two haplotypes were not detected in wild populations but were closely related to haplotypes found in the Rufiji River (one individual descended from one founder) and Shimoni, Kenya (two individuals descended from one founder) populations, suggesting nearby origins. Thus, the AZA-managed population of Angolan colobus likely originated from several localities, but all have mtDNA lineages associated with the subspecies C. a. palliatus, a Vulnerable subspecies. Examining founders' mtDNA haplotypes may be a useful addition to the zoo population management toolkit to help improve breeding recommendations by identifying individuals with rare haplotypes and revealing likely kinship among founders.

Non-Reproductive Sexual Behavior in Wild White-Thighed Colobus Monkeys (Colobus vellerosus).

Rare behaviors are often missing from published papers, hampering phylogenetic analyses. Here, we report, for the first time, masturbation and same-sex sexual behavior (SSB) in both male and female black-and-white colobus monkeys. We recorded these behaviors during 32months of observation (1573h of focal animal sampling) on Colobus vellerosus collected at the Boabeng-Fiema Monkey Sanctuary in Ghana. Males were observed masturbating and involved in SSB more than females. Subadult males were the age-sex class that engaged in both of these behaviors most often and a third of all SSB observed in young males occurred when they were forming an all-male band (AMB), which are temporally transient social groups in this species. Our data support that masturbation in males may be a sexual outlet for individuals that do not have a current sexual partner, while in females it may function in mate attraction by advertising receptivity. SSB may occur as an evolutionary byproduct but given the temporal clustering of observed events in males prior to AMB formation, our data best support the hypothesis that these behaviors facilitate male-male bonding (i.e., act as social glue). Within AMB's, males engage in coalitionary behavior to take over social groups containing females and strong bonds are important for success and later access to females, which could have selected for SSB in C. vellerosus.

DNA integrity and viability of testicular cells from diverse wild species after slow freezing or vitrification.

Cryopreservation of testicular tissues offers new possibilities to protect endangered species, genetically valuable individuals or even the fertility potential of prepubertal individuals who have died unexpectedly. However, the use of this technique still remains a challenge. In this study, slow freezing and vitrification of testicular tissue was investigated to find out which cryopreservation method could better preserve the viability and DNA integrity of testicular germ cells in diverse wild species. Testes were obtained post-mortem from 18 artiodactyls (wild boar, roe deer, dwarf goat, mhor gazelle, European mouflon, African forest buffalo, Malayan tapir, dorcas gazelle, Iberian ibex, gnu, red river hog), 5 primates (colobus monkey, capuchin monkey, mandrill), 8 carnivores (gray wolf, Persian leopard, binturong, European mink, American black bear, suricata), and 2 rodents (Patagonian mara). The testicles belonged to adult individuals and were cut into small pieces and cryopreserved by needle immersed vitrification or uncontrolled slow freezing using a passive cooling device. After warming or thawing, testicular tissues were enzymatically digested and two germ cell types were differentiated based on their morphology: rounded cells (spermatogonia, spermatocytes, and early spermatids) and elongated cells (elongated spermatids and spermatozoa). Cell viability was assessed by SYBR-14/propidium iodide while DNA fragmentation by TUNEL assay with fluorescence microscope. Our preliminary results revealed that our uncontrolled slow freezing method better preserved the viability and DNA integrity of elongated cells than vitrification. Such trend was observed in all species, being significant in artiodactyls, carnivores, and primates. Similarly, the viability and DNA integrity of rounded cells was also better maintained in primates by uncontrolled slow freezing, while in carnivores, vitrification by needle immersion showed better results in this type of cells. In artiodactyls and rodents both techniques preserved the viability of rounded cells in a similar manner, although the DNA integrity of these cells was greater after needle immersed vitrification in artiodactyls. In conclusion, the effectiveness of each cryopreservation method is affected by the phylogenetic diversity between species and cell type.

Assessing stress in wild black-and-white colobus monkeys non-invasively

Analysis of glucocorticoid profiles serves as a valuable, multi-faceted tool for insight into the behavior and physiology of wild populations. Recently, the measurement of fecal glucocorticoid metabolites (FCMs) has exploded in popularity due to its compatibility with noninvasive techniques and remote environments A critical first step is to perform a biological validation to ensure that the assay accurately reflect changes in FCM levels. We use an enzyme immunoassay (EIA) to perform a biological validation on samples collected from two males and six females in a wild population of Colobus vellerosus in response to three naturally occurring potential stressors. We also describe the FCM response pattern in the week following parturition in three females and examine the influence of sex, reproductive state, and time of day on the concentrations of baseline samples collected daily from 13 adult individuals over a period of four months. We validated the assay: FCM levels increase in response to natural stressors with a two-day lag. In the two days surrounding parturition, FMC levels increased. Baseline concentrations were affected by collection time and female reproductive state, with lactating females having lower concentrations than pregnant or cycling females. Thus, we successfully carried out the first validation and characterization of FCMs in a wild African colobine. This will serve as an essential foundation for future studies of C. vellerosus and similar wild primates whose objective is to investigate the role glucocorticoids play in responses to social and ecological challenges.

Growing population of the critically endangered white-thighed colobus monkey (Colobus vellerosus) from forest fragments in Ghana

Abstract The population of critically endangered white-thighed colobus monkeys (Colobus vellerosus) at Boabeng-Fiema Monkey Sanctuary (BFMS) is possibly the only growing population of this species in West Africa. We assessed the current population status of C. vellerosus in BFMS and the surrounding fragments in Ghana. We undertook a complete count of the population in 2020, and this data was combined with previously conducted complete counts from 1990 to 2014. Results show that the total population growth rate of colobus monkeys at BFMS and the surrounding forest fragments was 353.9% between the 1990 and 2020 censuses (at a rate of 11.8% annually). In the BFMS alone, the total population growth rate was 252.3% between 1990 and 2020 (i.e., at a rate of 8.4% annually). The total population growth rate in the surrounding forest fragments was 97.0% between the first census year of 1997 and the 2020 census (i.e., at a rate of 4.2% annually). The mean group size in the BFMS was 16.7 individuals (SD = 4.0; range = 9–25), while that of the surrounding forest fragments was 14.4 individuals (SD = 4.6; range = 9–23). The overall mean group size was 16.1 individuals (SD = 4.3; range = 9–25). An approximate ratio of one adult male to three adult females (1:3.4) and one adult female to one immature (1:1.2) is an indication that the population of C. vellerosus still has the potential to increase further when new suitable forest fragments are explored in the future. C. vellerosus has the potential to increase further in population in small, suitable fragments if habitat destruction and settlement expansion are managed with primate conservation intentions.

Attitudes of people living near Rau Forest Reserve, Tanzania towards Mount Kilimanjaro black and white colobus monkeys (Colobus caudatus)

Attitudes of people living near Rau Forest Reserve, Tanzania towards Mount Kilimanjaro black and white colobus monkeys (<i>Colobus caudatus</i>)

Population Status, Diurnal Activity Pattern, Feeding Ecology, and Habitat Association of Colobus Monkey (Colobus guereza) in Saja Forest, Kaffa Zone, Southwest Ethiopia

Colobus monkeys (Colobus guereza) are endemic to the Ethiopian plateau, distributed in different ecological habitats such as moist and deciduous forests, savanna woodlands, and montane forests. The population status, diurnal activity pattern, feeding ecology, and habitat association of Colobus guereza were investigated in Saja Forest, southwest Ethiopia, from June 2019 to February 2020, covering both wet and dry seasons. A total of 39 different types of transect were systematically established, distributed in both dense forest and in shrubland. Data were collected for 60 days in total across a wet and a dry season, both at dawn and in the afternoon. A total of 246 ± 39.1 individuals were recorded, of which 132 ± 24.4 and 114 ± 14.7 individuals were recorded during the wet and dry seasons, respectively. The population and group sizes did not statistically differ between the wet and dry seasons but shrubland habitat had fewer individuals and smaller groups than forest. Out of the recorded Colobus guereza, 33.5% were adult males, 34.5% were adult females, 28% were sub-adult males, 14.5% were sub-adult females, and 12.5% were juveniles/young. Feeding (29.5%) and resting (19.5%) were the most recorded daily activities for Colobus guereza. Young leaves were the largest (31%) contributor to the diet followed by mature leaves (22%) in both seasons. Other common dietary items were shoots (20%), barks (13%), fruits (11%), and flowers (3%). Colobus guereza were observed feeding on a diverse diet of 26 plant species belonging to 21 genera within 21 families. The habitats of these primates are currently diminishing due to anthropogenic activities such as agricultural expansion, human settlement, livestock grazing, and other forms of human wildlife conflict. Furthermore, guerezas are hunted for their skin and are also major sources of meat for the Menja people in the study area. Therefore, awareness creation for local people towards wildlife conservation is needed.

Distribution and diversity of primates and threats to their survival in the Awi Zone, northwestern Ethiopia.

Habitat loss and fragmentation affect the diversity and distribution of primates in a human-modified landscape. Ethiopia has a high diversity of primates, but increasing human pressure has negatively impacted their distribution and abundance across the country, primarily due to deforestation. To date, the diversity and distribution of primate species are poorly known in northwestern Ethiopia. From October 2020 until September 2021, we assessed the diversity and distribution of primate species in 26 forest patches in the Awi Zone, Northwestern Ethiopiausing line transect surveys, and weexamined the potential conservation threats to thesurvival of these taxa. Across transects, we encountered 459 groups of four primate taxa: olive baboons (Papio anubis), grivet monkeys (Chlorocebus aethiops), Boutourlini's blue monkeys (Cercopithecus mitis boutourlinii), and black-and-white colobus monkeys (Colobus guereza spp. guereza). The latter two are endemic to Ethiopia. We observed black-and-white colobus monkeys in all surveyed forest patches, while we observed Boutourlini's blue monkeys in 18 patches. Black-and-white colobus monkeys were the most frequently observed (n = 325 sighting; relative encounter frequency = 70.8%), while grivet monkeys (Chlorocebus aethiops) were the least (n = 34 sighting; relative encounter frequency = 7.4%) in the region. Similarly, the relative encounter frequency of olive baboons was 9.2% (n = 42 sighting).The overall mean group size for each species was: Boutourlini's blue monkeys (26.1 individuals), black-and-white colobus monkeys (8.8 individuals), grivet monkeys (34.1 individuals), and olive baboons (41.4 individuals). We identified agricultural expansions, exotic tree plantations, deforestations, firewood collections, livestock grazing, and killings over their crop-feeding behaviors as the main threats to primates and their habitats in the region. This study provides crucial information on an area likely to support primate speciesthat we know very little about. Assigning protected connecting forest patches should be an urgent priority for the conservation of the primates in this region.

Chimpanzee (Pan troglodytes) gaze is conspicuous at ecologically-relevant distances

Chimpanzee (Pan troglodytes) sclera appear much darker than the white sclera of human eyes, to such a degree that the direction of chimpanzee gaze may be concealed from conspecifics. Recent debate surrounding this topic has produced mixed results, with some evidence suggesting that (1) primate gaze is indeed concealed from their conspecifics, and (2) gaze colouration is among the suite of traits that distinguish uniquely social and cooperative humans from other primates (the cooperative eye hypothesis). Using a visual modelling approach that properly accounts for specific-specific vision, we reexamined this topic to estimate the extent to which chimpanzee eye coloration is discriminable. We photographed the faces of captive chimpanzees and quantified the discriminability of their pupil, iris, sclera, and surrounding skin. We considered biases of cameras, lighting conditions, and commercial photography software along with primate visual acuity, colour sensitivity, and discrimination ability. Our visual modeling of chimpanzee eye coloration suggests that chimpanzee gaze is visible to conspecifics at a range of distances (within approximately 10 m) appropriate for many species-typical behaviours. We also found that chimpanzee gaze is discriminable to the visual system of primates that chimpanzees prey upon, Colobus monkeys. Chimpanzee sclera colour does not effectively conceal gaze, and we discuss this result with regard to the cooperative eye hypothesis, the evolution of primate eye colouration, and methodological best practices for future primate visual ecology research.

Do zoo-housed primates retreat from crowds? A simple study of five primate species.

An animal's welfare state is directly influenced by the mental state, which is shaped by experiences within the environment throughout the animal's life. For zoo-housed animals, visitors to the zoo are a large part of that environment and a fluctuating influence within it. This study examines the impact of zoo visitors on the space use of five species of zoo-housed primates (Eastern black-and-white colobus monkeys, Colobus guereza, n = 5, Allen's swamp monkeys, Allenopithecus nigroviridis, n = 2, DeBrazza's monkeys, Cercopithecus neglectus, n = 3, Bolivian gray titi monkeys, Callicebus donacophilus, n = 3, and crowned lemurs, Eulemur coronatus, n = 3). Specifically, we considered whether primates' distance from visitor areas changed as crowd sizes increased. Data were collected using the ZooMonitor app. Observers recorded spatial coordinates for each animal over periods ranging from 12 to 32 months. Data were analyzed using two types of regression models (linear and logistic) to examine the influence of visitors on the location of the primates. Both analyses revealed a statistically significant but small decrease in primate distance from visitor viewing glass as the number of visitors increased. Behavioral indicators of welfare were also unaffected by the presence of visitors. These results suggest that, with additional validation, distance from visitors may be one promising, simple way to evaluate the influence of visitors on primate welfare.

Mitochondrial genetic diversity and divergence dating of Angolan colobus monkeys (Colobus angolensis) in the eastern forests of Kenya and Tanzania: Implications for subspeciation and reconstructing historical biogeography.

Whether the Colobus angolensis that reside in the fragmented forests in eastern Kenya and Tanzania represent one subspecies or two has been debated for 50 years. Morphological and more recent genetic and ecological studies suggest that these populations represent two subspecies, C. a. palliatus and C. a. sharpei. However, their distribution of mitochondrial variation remains unresolved since the genetic study only characterized four populations at the range ends. Therefore, we characterized five populations in the area of the hypothesized subspecies divide. We identified eight new haplotypes which, combined with those previously identified, provided 26 haplotypes from nine populations for analysis. Haplotypes found south of the Rufiji River cluster together but separately from northern haplotypes. The largest sequence differences within cytochrome b occur between population pairs representing opposite sides of the river; their mean difference (1.5%) is more than that of other primate subspecies. Analysis of molecular variance attributes most of the variation to that north versus south of the river. These results support the previous subspecies distinction between C. a. palliatus (northern) and C. a. sharpei (southern), divided by the Rufiji River. The estimated time of the most recent common ancestor of all haplotypes indicates that the subspecies have been isolated from each other for approximately 550,000 years. The common ancestor of northern and southern haplogroups was 370,000 and 290,000 years ago, respectively. Nevertheless, the correlation between genetic and geographic distances suggests that isolation-by-distance contributed to population structuring. Significant variation among populations, with only three haplotypes shared between populations, also indicates that an extended period of isolation drove population distinctiveness. Considering these results, we evaluate hypotheses about the founding and differentiation of these subspecies during Pleistocene climatic fluctuations and propose a novel, more direct migration route from Central Africa to their current range navigating Lake Tanganyika, the central Tanzanian corridor, and the Rufiji River.