Jean-Pierre Gautier

A primate radiation : evolutionary biology of the African guenons

Providing a synthesis of our present knowledge of a colourful group of monkeys - the African Guenons, this book examines some of the current views on mammalian community structure and speciation, as well as on the phylogeny of African primates.

Diets of two sympatric colobines in zaire : more evidence on seed-eating in forests on poor soils

Recent studies have shown that the so- called folivorous African colobines eat a significant amount of seeds. There is disagreement as to whether seed-eating is due to the poor quality of tree foliage, due in turn to poor soils, or to the fact that seeds are a normal part of colobine diets. To test these hypotheses, we studied feeding of red colobus monkeys, Colobus badius tholloni,and black-and-white colobus monkeys, Colobus angolensis angolensis,in a tropical rain forest of Central Zaire (Salonga National Park). We conducted studies on soil properties, vegetation composition, and the availability and chemical contents of food plants simultaneously. Soils were very acid, with a high percentage of sand, very low cation exchange capacity, and very low exchangeable bases. The forest was dominated by legumes (45.6% of trees), among which the Caesalpinioideae were the best represented (85%). C. badiusfed mostly on leaves (61%) and seeds (33%), legumes making up 65% of their diet. C. angolensisfed mostly on seeds (50%) and leaves (27%); 39% of their diet came from legumes. The two species tended to select items richer in crude protein or lipids or both. Total phenolics and condensed tannins were abundant in the foliage and seeds but were poor predictors of colobine choice of food. Intersite comparisons show that colobines in Zaire ate a higher proportion of seeds than all other related species so far studied in Africa and that the Salonga forest had among the poorest soils and harbored the highest percentage of Caesalpinioideae. Our results confirm that seed- eating is more common among colobines living in areas where soils are poor. They strongly suggest that this link is mediated through forest composition, especially the abundance of legumes, and that the development of seed- eating results both from the high availability of nutrient-rich seeds and from the poor quality of mature tree foliage.

Primate diet and biomass in relation to vegetation composition and fruiting phenology in a rain forest in Gabon

To test the hypothesis that primate populations are limited by food resources, we studied the feeding ecology of three cercopithecines and one colobine in a rain forest in central Gabon. Simultaneously, we monitored the fruiting phenology of trees and estimated the biomass of the monkey community. The Makandé Forest is dominated by Caesalpiniaceae and characterized by a lack of secondary vegetation and of trees species producing fleshy fruits. Fruit production was irregular intra- and interannually. Fruiting peaks of dry fruits (mainly Caesalpiniaceae) and of fleshy fruits occurred at the same period. However, interseasonal and interannual variability was greater in Caesalpiniaceae than in other families. As a result, the Makandé forest is subject to bottlenecks when food is scarce. On an annual basis, seeds (primarily Caesalpiniaceae) dominated the diet of all monkeys. On a seasonal basis, cercopithecines preferentially consumed fleshy fruits as long as they were available, whereas colobines increased consumption of young leaves when seed availability declined. The consumption of mature leaves was low. The monkey community biomass (ca. 204 kg/km2) is one of the lowest in Central Africa. We suggest that both cercopithecine and colobine populations are limited as a result of the combined effect of the dominance of Caesalpiniaceae, which provide dry fruits according to a mast-fruiting pattern and mature leaves of low quality, and the lack of seral successional stages, which provide fleshy fruit on a more regular pattern and leaves of better quality. During the period of food scarcity, cercopithecines should suffer from the low availability of fleshy fruit, which are their favorite food. At the same period, colobines should be limited by the low availability of edible leaves. Similar low primate biomasses are found in forests dominated by Caesalpiniaceae or Lecythidaceae in South America and in Dipterocarpaceae forests in South Asia, which suggests that their biological characteristics, in particular dry fruits and mast fruiting, are unfavorable to monkey populations. Our results confirm that habitat mosaics may support larger populations of primary consumers than homogeneous primary forests can.

Seed dispersal versus seed predation: an inter-site comparison of two related African monkeys

C. pogonias and C. wolfi plant diets were studied in two sites, in Gabon and Zaire and compared with fruit availabilities. Monkeys in Gabon were found to be mainly fruit pulp-eaters while monkeys in Zaire were alternately seed-eaters, aril-eaters or leaf-eaters. These differences were related to differences in the availability of fruit categories: fleshy fruits were found to be much more abundant in Gabon than in Zaire forests. As a result, monkeys in Gabon were found to be mainly seed-dispersers while monkeys in Zaire were found, to a large extent, to be seed-predators. Results are discussed in terms of phenotypic flexibility in monkey feeding behavior, diversity of plant-monkey interactions, geographic variability of keystone plant resources, and their implications for forest management practices. The low availability of fleshy fruit species in Zaire is hypothezized to result from the poor soil conditions.

On the Occurrence of Nocturnal and Diurnal Loud Calls, Differing in Structure and Duration, in Red Howlers (Alouatta seniculus) of French Guyana

Structural patterns and 24-hour rhythms of the loud calls of Alouatta seniculus were studied in a wild population living in primary rain forest in French Guyana. These loud calls are low-pitched noisy hoarse sounds with an upper frequency limit around 3,000 Hz. Their spectral energy is concentrated in two clusters, each of which includes two dominant frequency bands. Loud calls comprise one or several vocal units with alternating crescendos and decrescendos. Five different types of units can be distinguished. Depending on their duration, loud calls can be roughly divided into two classes: long and short calls. Long calls (mean duration 3.28 min) include three phases. After the end of the calls, one or two blowing noises of weak amplitude may be heard. Short calls (mean duration 11 s) are frequently uttered in clusters of several calls given by the same troop at intervals of a few seconds to a few minutes. Observations carried out over the 24-hour cycle show that the calling rate is higher at night, with a peak around dawn and that long calls are more frequent at night, while short calls are mainly heard during the day. Most long calls are emitted from the sleeping sites and carry great distances. They probably play a role in inter-troop communication. Despite their intensity, short calls are used for short-distance communication. The occurrence of two different loud calls in Guyanan howlers and the amount of night calling in these monkeys are discussed with reference to studies of other howler populations.

The distribution ofPapio hamadryas in Saudi Arabia: Ecological correlates and human influence

We describe the distribution ofPapio hamadryas in Saudi Arabia in relation to ecological constraints and recent changes in human activities and land use. Baboons are present only in the Sarawat and southern Hijaz mountains, which border the Red Sea in the southwestern region of the country. Our distribution data confirms that their range, described as of 1981, should be extended 200 km northward, from 21°40N (Taif) to 23°20N (Al Akhal). This new limit of range is not due to recent deployment of the baboons, but instead corrects inaccurate surveys by previous reporters. Ecological factors that affect baboon distribution include low annual rainfall (less than 100 mm per yr), which limits baboon range in the north and east, and the absence of suitable sleeping cliffs in the coastal plain, which limits their range in the west. Relative abundance is discussed in relation to vegetation and predator distribution. Human influence is also significant: commensal troops persist near urban centers, while agricultural development reduces baboon numbers in some regions. Habitat characteristics are discussed in comparison with similar data on the range of hamadryas in Ethiopia.

Group composition in wild and commensal hamadryas baboons: A comparative study in Saudi Arabia

Papio hamadryas was surveyed throughout its range in Saudi Arabia and was observed at altitudes ranging from 0 to 2300 m. Wild populations occur along the whole range of altitude, while commensal populations are only found above 850 m altitude. No variation in group size was found with altitude. Comparison of wild and commensal populations showed the following. (1) Their composition in terms of age and sex classes, overall adult sex ratios, and group size does not significantly differ. (2) Groups of both populations include, in similar proportions, three types of parties: one-male units (>70%), two-male units (>13%), and a few other units of variable composition. (3) The mean size of commensal parties is significantly larger than in the wild population; specifically one-male units are larger in the commensal population due to a larger number of females per male. Thus, female distribution in commensal groups is more inequitable than that in wild groups. (4) Finally, the number of females included in two-male units increases with altitude. These differences are discussed in terms of food availability and predator pressure and are compared with results obtained on other Arabian and Ethiopian populations.

Formal monkey linguistics

Abstract We argue that rich data gathered in experimental primatology in the last 40 years can benefit from analytical methods used in contemporary linguistics. Focusing on the syntactic and especially semantic side, we suggest that these methods could help clarify five questions: (i) what morphology and syntax, if any, do monkey calls have? (ii) what is the ‘lexical meaning’ of individual calls? (iii) how are the meanings of individual calls combined? (iv) how do calls or call sequences compete with each other when several are appropriate in a given situation? (v) how did the form and meaning of calls evolve? We address these questions in five case studies pertaining to cercopithecines (Putty-nosed monkeys, Blue monkeys, and Campbell’s monkeys), colobinae (Guereza monkeys and King Colobus monkeys), and New World monkeys (Titi monkeys). The morphology mostly involves simple calls, but in at least one case (Campbell’s -oo) we find a root-suffix structure, possibly with a compositional semantics. The syntax is in all clear cases simple and finite-state. With respect to meaning, nearly all cases of call concatenation can be analyzed as conjunction. But a key question concerns the division of labor between semantics, pragmatics and the environmental context (‘world’ knowledge and context change). An apparent case of dialectal variation in the semantics (Campbell’s krak) can arguably be analyzed away if one posits sufficiently powerful mechanisms of competition among calls, akin to scalar implicatures. An apparent case of non-compositionality (Putty-nosed pyow-hack sequences) can be analyzed away if one further posits a pragmatic principle of ‘urgency’, whereby threat-related calls must come early in sequences (another potential case of non-compositionality – Colobus snort-roar sequences – might justify assigning non-compositional meanings to complex calls, but results are tentative). Finally, rich Titi sequences in which two calls are re-arranged in complex ways so as to reflect information about both predator identity and location are argued not to involve a complex syntax/semantics interface, but rather a fine-grained interaction between simple call meanings and the environmental context. With respect to call evolution, we suggest that the remarkable preservation of call form and function over millions of years should make it possible to lay the groundwork for an evolutionary monkey linguistics, which we illustrate with cercopithecine booms, and with a comparative analysis of Blue monkey and Putty-nosed monkey repertoires. Throughout, we aim to compare possible theories rather than to fully adjudicate between them, and our claims are correspondingly modest. But we hope that our methods could lay the groundwork for a formal monkey linguistics combining data from primatology with formal techniques from linguistics (from which it does not follow that the calls under study share non-trivial properties, let alone an evolutionary history, with human language).

Additional Data on the Distribution of Cercopithecus (lhoesti) solatus

The sun-tailed monkey Cercopithecus (lhoesti) solatus Harrison 1988, an endemic guenon from Gabon [1], has an unfragmented distribution area estimated at about 10,300 km2 mainly centered in the Forêt des Abeilles, central Gabon [2]. It is limited in the north by the Ogooué river and in the east by the Ogooué, Lolo and Bouenguidi rivers. The southern limit is unclear, but Gautier et al. [2] suggested that it corresponds to an elevation in altitude because the sun-tailed monkey is absent at altitudes higher than 500 m above sea level (asl). In the west, the limit was considered to be the Offoué river, but it was suspected that the species could occur on the left bank of this river. This was confirmed by forestry prospector C. Wilks [pers. commun.] who saw C. solatus monkeys along the left bank of the Offoué river in the south of the Lopé Reserve in 1994, and by White and Mackanga [3] who observed individuals in the northern part of the Lopé Reserve, about 30 km west of the Offoué river. Thus, as recommended in the African Primates Conservation Action Plan [4], the western limit of the species needs to be clarified. During 1996, we carried out primate census surveys in the south of the Lopé Reserve, near the proposed western limit of C. solatus. New data are presented in this paper on the distribution of C. solatus and its ecological patterns are discussed.

Do black colobus in mixed-species groups benefit from increased foraging efficiency?

Seed-eating in the African colobines varies according to species and site. Because seeds are more palatable than leaves, seed-eating is considered to result from a positive choice by colobines. A year-long study of the diet of Colobus satanas in Central Gabon confirmed that this species eats a significant amount of seeds. Furthermore, despite large variations in fruit availability, this amount consistently increases when colobines feed in mixed-species troops including frugivorous cercopithecines and mangabeys. Our results show that colobines benefit when feeding in mixed troops and suggest that polyspecific associations are an adaptive strategy that could have positive effects on colobine populations, especially during the period of fruit shortage.