Cryptic diversity and species boundaries within the Paragalago zanzibaricus species complex.

The recently described genus Paragalago is a complex of several nocturnal and morphologically cryptic species distributed in the forests of eastern Africa. Species diversity within this genus has been mainly described using species-specific differences in their loud calls. However, molecular data are still lacking for this group and species boundaries remain unclear. In this study, we explore species diversity within the zanzibaricus-complex using a combination of mitochondrial and nuclear data and comparing multiple species delimitation methods. Our results consistently support the existence of three independent lineages, P. cocos, P. zanzibaricus, and P. granti, confirming previous hypotheses based on vocal data. We conclude that these three lineages represent valid cryptic species and we hypothesize that speciation within this complex was characterized by cycles of forest expansion and contraction in the Plio-Pleistocene.

Species Boundaries within Morphologically Cryptic Galagos: Evidence from Acoustic and Genetic Data.

Describing primate biodiversity is one of the main goals in primatology. Species are the fundamental unit of study in phylogeny, behaviour, ecology and conservation. Identifying species boundaries is particularly challenging for nocturnal taxa where only subtle morphological variation is present. Traditionally, vocal signals have been used to identify species within nocturnal primates: species-specific signals often play a critical role in mate recognition, and they can restrict gene flow with other species. However, little research has been conducted to test whether different "acoustic forms" also represent genetically distinct species. Here, we investigate species boundaries between two putative highly cryptic species of Eastern dwarf galagos (Paragalago cocosand P. zanzibaricus). We combined vocal and genetic data: molecular data included the complete mitochondrial cytochrome b gene (1,140 bp) for 50 samples across 11 localities in Kenya and Tanzania, while vocal data comprised 221 vocalisations recorded across 8 localities. Acoustic analyses showed a high level of correct assignation to the putative species (approx. 90%), while genetic analyses identified two separate clades at the mitochondrial level. We conclude that P. cocos and P. zanzibaricus represent two valid cryptic species that probably underwent speciation in the Late Pliocene while fragmented in isolated populations in the eastern forests.

Low Geographic and Subspecific Variation in the Loud Call of the Widespread and Phenotypically Cryptic Northern Lesser Galago (Galago senegalensis) Suggests Taxonomic Uniformity.

Like other nocturnal primates, many species of galago (Galagidae) are phenotypically cryptic, making their taxonomic status difficult to resolve. Recent taxonomic work has disentangled some of the confusion. This has resulted in an increase in the number of recognised galago species. The most widespread galago species, and indeed the most widespread nocturnal primate, is the northern lesser galago (Galago senegalensis) whose geographic range stretches >7,000 km across Africa. Based on morphology, 4 subspecies are currently recognised: G. s. senegalensis, G. s. braccatus, G. s. sotikae and G. s. dunni. We explore geographic and subspecific acoustic variation in G. senegalensis, testing three hypotheses: isolation by distance, genetic basis, and isolation by barrier. There is statistical support for isolation by distance for 2 of 4 call parameters (fundamental frequency and unit length). Geographic distance explains a moderate amount of the acoustic variation. Discriminant function analysis provides some degree of separation of geographic regions and subspecies, but the percentage of misdesignation is high. Despite having (putative) parapatric geographic ranges, the most pronounced acoustic differences are between G. s. senegalensis and G. s. dunni. The findings suggest that the Eastern Rift Valley and Niger River are significant barriers for G. senegalensis. The acoustic structures of the loud calls of 121 individuals from 28 widespread sites are not significantly different. Although this makes it unlikely that additional unrecognised species occur within G. senegalensis at the sites sampled, vast areas of the geographic range remain unsampled. We show that wide-ranging species do not necessarily exhibit large amounts of variation in their vocal repertoire. This pattern may also be present in nocturnal primates with smaller geographic ranges.

Sleep patterns, daytime predation, and the evolution of diurnal sleep site selection in lorisiforms.

OBJECTIVES: Synthesize information on sleep patterns, sleep site use, and daytime predation at sleep sites in lorisiforms of Asia and Africa (10 genera, 36 species), and infer patterns of evolution of sleep site selection. MATERIALS AND METHODS: We conducted fieldwork in 12 African and six Asian countries, collecting data on sleep sites, timing of sleep and predation during daytime. We obtained additional information from literature and through correspondence. Using a phylogenetic approach, we established ancestral states of sleep site selection in lorisiforms and traced their evolution. RESULTS: The ancestral lorisiform was a fur-clinger and used dense tangles and branches/forks as sleep sites. Use of tree holes and nests as sleep sites emerged ∼22 Mya (range 17-26 Mya) in Africa, and use of bamboo emerged ∼11 (7-14) Mya in Asia and later in Africa. Fur clinging and some sleep sites (e.g., tree holes, nests, but not bamboo or dense tangles) show strong phylogenetic signal. Nests are used by Galagoides, Paragalago, Galago and Otolemur; tree holes by Galago, Paragalago, Sciurocheirus and Perodicticus; tangles by Nycticebus, Loris, Galagoides, Galago, Euoticus, Otolemur, Perodicticus and Arctocebus; all but Sciurocheirus and Otolemur additionally sleep on branches/forks. Daytime predation may affect sleep site selection and sleep patterns in some species of Nycticebus, Galago, Galagoides, Otolemur and Perodicticus. Most lorisiforms enter their sleep sites around sunrise and leave around sunset; several are active during twilight or, briefly, during daytime. CONCLUSION: Variations in sleep behavior, sleep patterns and vulnerability to daytime predation provide a window into the variation that was present in sleep in early primates. Overall, lorisiforms use the daytime for sleeping and no species can be classified as cathemeral or polycyclic.

A giant among dwarfs: a new species of galago (Primates: Galagidae) from Angola.

OBJECTIVES: Based on vocalization recordings of an unknown galago species, our main objectives were to compare morphology and call structure with known closely-related taxa and describe a new species of galago. MATERIALS AND METHODS: We conducted field surveys in three forest habitats along the escarpment region in western Angola (Kumbira Forest, Bimbe Area, and Northern Scarp Forest), and examined galago specimens from museums worldwide. We digitized and analyzed calls using Avisoft SASLab Pro software. We also compared museum specimens from Angola with other Galago and Galagoides specimens, and conducted comparative analyses (ANOVA and between group principle component analysis) based on a set of twelve linear measurements of skulls and teeth. RESULTS: We describe the new species to which we give the name Angolan dwarf galago, Galagoides kumbirensis sp. nov. The new species has a loud and characteristic crescendo call, used by other Galagoides spp. (sensu stricto) in West Africa to attract companions and repel rivals. However, this call shows species-typical differences from its closest relatives. Galagoides kumbirensis sp. nov. is also distinguished by differences in the skull morphology, pelage color and facial markings, as well as a larger body size, similar to that of Galago moholi, which is not known to be sympatric. CONCLUSION: This discovery points to the importance of Angolan forests as refuges for endemic biodiversity. These forests are under severe threat from overexploitation, and there is an urgent need to establish conservation measures and designate protected areas.

Niche partitioning and environmental factors affecting abundance of strepsirrhines in Angola.

The African nocturnal primates (galagos, pottos, and angwantibos: suborder Strepsirrhini) are the result of the first major primate radiation event in Africa, and are found in different primate communities spread across the entire sub-Saharan Africa. Thus, they represent an interesting group of taxa to investigate community strategies to avoid interspecific competition. Here, we present the result of the first study on nocturnal primate communities in western Angola. We aimed to identify habitat factors influencing strepsirrhine abundance, collect evidence of spatial niche segregation, and discuss possible indications of competitive exclusion in this region. We conducted nocturnal surveys at four study sites: Kumbira, Bimbe, Northern Scarp, and Calandula. At each encounter we recorded species, group size, height of animals above ground, and GPS location. We sampled vegetation using the point-centered quarter method and collected data on canopy cover, disturbance, and undergrowth density. We observed a total of five strepsirrhine species with varying community structures. We did not encounter Galagoides thomasi but we recorded a new species Galagoides sp. nov. 4. Levels of disturbance, canopy cover and undergrowth density were the habitat factors that most influenced variation in abundance of Galagoides demidovii and Perodicticus edwardsi, the latter also preferring the habitat with higher tree density. Vertical separation between sympatric strepsirrhines was strongest in Northern Scarp, where overall relative abundance was also highest. Competitive exclusion between G. thomasi and G. sp. nov. 4 may explain why the former was not present within the Angolan Escarpment sites. We observed coexistence between mainly allopatric Otolemur crassicaudatus and P. edwardsi in Kumbira, and of Galago moholi and G. demidovii in Calandula. Both unusual combinations showed some levels of spatial segregation. Habitat characteristics of the Escarpment region are likely to allow for unique nocturnal primate species assemblages. We urge immediate conservation interventions in the Angolan Escarpment.

Density estimates of Panamanian owl monkeys (Aotus zonalis) in three habitat types.

The resolution of the ambiguity surrounding the taxonomy of Aotus means data on newly classified species are urgently needed for conservation efforts. We conducted a study on the Panamanian owl monkey (Aotus zonalis) between May and July 2008 at three localities in Chagres National Park, located east of the Panama Canal, using the line transect method to quantify abundance and distribution. Vegetation surveys were also conducted to provide a baseline quantification of the three habitat types. We observed 33 individuals within 16 groups in two out of the three sites. Population density was highest in Campo Chagres with 19.7 individuals/km(2) and intermediate densities of 14.3 individuals/km(2) were observed at Cerro Azul. In la Llana A. zonalis was not found to be present. The presence of A. zonalis in Chagres National Park, albeit at seemingly low abundance, is encouraging. A longer-term study will be necessary to validate the further abundance estimates gained in this pilot study in order to make conservation policy decisions.

Remarkable ancient divergences amongst neglected lorisiform primates.

Lorisiform primates (Primates: Strepsirrhini: Lorisiformes) represent almost 10% of the living primate species and are widely distributed in sub-Saharan Africa and South/South-East Asia; however, their taxonomy, evolutionary history, and biogeography are still poorly understood. In this study we report the largest molecular phylogeny in terms of the number of represented taxa. We sequenced the complete mitochondrial cytochrome b gene for 86 lorisiform specimens, including ∼80% of all the species currently recognized. Our results support the monophyly of the Galagidae, but a common ancestry of the Lorisinae and Perodicticinae (family Lorisidae) was not recovered. These three lineages have early origins, with the Galagidae and the Lorisinae diverging in the Oligocene at about 30 Mya and the Perodicticinae emerging in the early Miocene. Our mitochondrial phylogeny agrees with recent studies based on nuclear data, and supports Euoticus as the oldest galagid lineage and the polyphyletic status of Galagoides. Moreover, we have elucidated phylogenetic relationships for several species never included before in a molecular phylogeny. The results obtained in this study suggest that lorisiform diversity remains substantially underestimated and that previously unnoticed cryptic diversity might be present within many lineages, thus urgently requiring a comprehensive taxonomic revision of this primate group.

Social organization of the Milne-Edward's potto.

Nocturnal prosimian primates are often seen alone during their nocturnal activities, and are therefore categorized as "solitary". Recent research has shown that these animals actually possess social networks that differ among species. Here we present new information on one of the lesser studied prosimian primates, the potto, derived from fieldwork in Cameroon that employed radiotelemetry and behavioral observations. An analysis of association patterns and home-range overlaps between animals revealed that pottos associated more frequently with conspecifics than expected for a supposedly solitary primate. Certain males and females that were seen together regularly throughout the study, but were not observed to have contact with other pottos of the opposite sex, were labeled as "pairs". These pairs were found to have higher levels of association with each other than with other conspecifics. The only affiliative and sexual behaviors observed between members of the opposite sex occurred within the pairs of pottos. Although given the secretive nature of these primates, the possibility of extrapair copulations cannot be ruled out, their relatively small testis size indicates that sperm competition is not so important for this species.

Species-typical patterns of infant contact, sleeping site use and social cohesion among nocturnal primates in Africa.

This paper provides a checklist and summary of what is currently known of the variation in infant contact, sleeping site preference and aspects of social cohesion in the nocturnal primates of Africa. Genera and species are compared, based on previously unpublished field observations and a review of the literature. There is a clear pattern of similarity between the species within each genus and distinct differences between genera. Species in the same genus tend to be ecologically equivalent and replace each other allopatrically, whereas species in different genera are more likely to be sympatric, with up to 6 species living together. Maximum sympatry within genera is found in Otolemur and Galagoides, where species are ecologically divergent. This may reflect an ancient origin of species within these genera or suggest that further taxonomic revision is required at the generic level. Some data are recorded for the first time for species that have only recently been separated (cryptic species), but some taxa remain very poorly known. It is concluded that field studies are still at an elementary stage and further research with radio tracking is urgently needed in the face of rapidly declining habitats.