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Stereotypical vocalisations can facilitate long-distance communication in dense and, thus, sound-degrading forest habitats. Despite this, primate vocal repertoires often also include gradations between different call types that are used in social interactions. Because many nocturnal primates show a solitary social structure, it has been difficult to assess the role vocalisations play in mediating their social encounters. Here we aim to expand on the call types of Sahamalaza sportive lemurs, Lepilemur sahamalaza, as well as investigate their variability and use in social contexts. Through long-term behavioural observations with concurrent recording of vocalisations of known and unknown individuals between 2015 and 2016, we described three previously unrecorded calls and expanded the vocal repertoire to 12 different call types that were used in both targeted social and broadcast solitary contexts. Linear discriminant function and cluster analysis supported initial observations that vocalisations within call types were variable and that at least two call types can be classified as “graded,” contrary to expectations. Despite variations in call-context connections, no clear patterns of call use could be established. However, measurable differences in vocal behaviour between the seasons and the sexes indicate that calling is used in a reproductive context, similar to other nocturnal and diurnal primates.

In response to predation pressure by raptors, snakes, and carnivores, primates employ anti-predator behaviours such as avoiding areas of high predation risk, cryptic behaviour and camouflage, vigilance and group formation (including mixedspecies associations), and eavesdropping on other species’ alarm calls. After detecting a predator, primates can produce alarm calls, show predator-specific escape strategies or even mob the predator. It remains unclear how solitary nocturnal primates respond to diurnal predation pressure while they sleep or rest. The aim of this study was to investigate the diurnal anti-predator behaviour of the nocturnal and solitary Sahamalaza sportive lemur, Lepilemur sahamalazensis, which regularly rests in exposed locations. We observed the responses of 32 Sahamalaza sportive lemurs to playbacks of territorial calls of an aerial predator (Madagascar harrier hawk), mating calls of a terrestrial predator (fossa), and the contact calls of a medium-sized bird (crested coua) as a control, at different diurnal sleeping sites. Lemurs never showed a flight response after replays of predator or control calls, but regularly froze after harrier hawk calls. Lemurs scanned the sky immediately after playback of harrier hawk calls, and the ground or trees after fossa calls. Lemur vigilance increased significantly after both predator calls. After crested coua calls the animals became significantly less vigilant, suggesting that contact calls of this bird serve as indicators of predator absence. We found no response differences between different types of sleeping sites. Our results show that resting Sahamalaza sportive lemurs recognise predator vocalisations as indicators of increased predation risk, discern vocalizations of different predators, and employ anti-predator behaviours specific for different predator classes. Their behavioural responses while resting or sleeping are comparable to those of active primates, and their response rate of 80% shows that this solitary and nocturnal primate is constantly aware of its environment.

Abstract

We provide an anecdote of daytime activity within the northern giant mouse lemur (Mirza zaza), a small threatened primate that has always been considered strictly nocturnal, in the Anabohazo forest of northwestern Madagascar. During the dry season, we witnessed two individual M. zaza travelling separately in the afternoon period of three different days. Our observations indicate that M. zaza may undertake activity that is essential for their survival within both the light and dark periods of a day-cycle, and our findings suggest that the activity cycle of this species may be highly flexible. These observations have important implications to understand the evolution of activity patterns in M. zaza, and these findings warrant future, long term studies to establish the annual activity patterns of M. zaza and to determine whether this species is potentially cathemeral and why.

Prior herpetological surveys in 1996 and 2000 identified 14 species of amphibians and 32 species of reptiles from the Sahamalaza Peninsula. This work increases the total number of amphibian and reptile species known from this area to 20 and 43 respectively. To maximise our chances of species detection, survey effort covered the entire wet season and part of the dry season, and utilised a combination of opportunistic searching, transect searching, pitfall trapping, and acoustic recording. We identified species through an integrative taxonomic approach, combining morphological, bioacoustic and molecular taxonomy. Together, this enabled the detection of cryptic and seasonally inactive species that were missed in the shorter prior surveys that relied on morphological identification alone. The taxonomic identification of amphibians utilised a fragment of the mitochondrial 16S rRNA gene; taxonomic identification of reptiles utilised a fragment of the mitochondrial COI gene, and when necessary, also mitochondrial fragments of the 16S rRNA ND1, ND2, ND4 genes. All sequences were deposited in Genbank and COI sequences were also deposited in the BOLD database to foster taxonomic identification of malagasy reptiles. We report two new taxa: a species of Boophis, since described as B. ankarafensis, and a candidate new species of microhylid (genus: Stumpffia). We document range expansions of Boophis tsilomaro, Cophyla berara, Blaesodactylus ambonihazo beyond their type localities. Along with significant range expansions across a range of taxa, including Blommersia sp. Ca05, Boophys brachychir, Brookesia minima, Ebenavia inunguis, Geckolepis humbloti, Madascincus stumpffi, Pelomedus subrufa and Phelsuma kochi. Forest in the peninsula is under extreme pressure from human exploitation. Unless unsustainable agricultural and pastoral practices encroaching on these habitats halt immediately, both forest and the species that occur there, several of which appear to be local endemics, may be irreversibly lost.