GROUP STRUCTURE AND ACTIVITY RHYTHM IN LEMUR MONGOZ (PRIMATES, LEMURIFORMES) ON AND MOHELI ISLANDS, COMORO ARCHIPELAGO

IAN TATTERSALL

VOLUME 53: PART 4 ANTHROPOLOGICAL PAPERS OF THE AMERICAN MUSEUM OF NATURAL HISTORY NEW YORK :1976 4 .4.4.4.4 .4.4

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11 I, -_7 Tf., GROUP STRUCTURE AND ACTIVITY RHYTHM IN LEMUR MONGOZ (PRIMATES, LEMURIFORMES) ON ANJOUAN AND MOHELI ISLANDS, COMORO ARCHIPELAGO

IAN TATTERSALL Associate Curator, Department ofAnthropology TheAmerncan Museum ofNatural History

VOLUME 53: PART 4 ANTHROPOLOGICAL PAPERS OF THE AMERICAN MUSEUM OF NATURAL HISTORY NEW YORK: 1976 ANTHROPOLOGICAL PAPERS OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 53, Part 4, pages 367-380, figures 1-6, tables 1, 2

Issued December 30, 1976 Price. $1.1 5

ISSN 0065-9452

This Part completes Volume 53.

Copyright i The American Museum of Natural History 1976 ABSTRACT A previous study in Madagascar revealed the day. This difference appears to be environ- Lemur mongoz to be nocturnal and to exhibit mentally linked. Further, on Anjouan L. mongoz pair-bonding. Subsequent work in the Comoro Is- exhibits pair-bonding and the formation of lands has shown that, whereas in the warm, "family" groups, but on Moheli there is variation seasonal lowland areas of Moheli and Anjouan L. in group structure. It is possible that group com- mongoz is likewise nocturnal, in the humid high- position of L. mongoz on Moheli undergoes lands of Anjouan these animals are active during seasonal change.

RItSUMI! Une premiere etude de Lemur mongoz de jour. Cette difference semble etre liee a Madagascar a revele que celui-ci est nocturne et l'environnement. De plus, a Anjouan Lemur qu'il vit par couples de deux adultes avec ou sans mongoz se montre en couples de deux adultes enfants de moins de deux ans. Une etude sui- avec ou sans enfants, alors qu'a Moheli on ren- vante faite aux Comores a montre que dans les contre une certaine variete dans la structure du basses terres chaudes et saisonnieres de Moheli et groupe. I1 est possible que la composition de d'Anjouan Lemur mongoz est egalement noc- groupe de Lemur mongoz a Moheli soit sujette a turne, alors que dans les hauteurs humides des changements saisonniers. d'Anjouan ces animaux sont actifs pendant le

INTRODUCTION With the exception of Lemur fulvus, the only Tattersall, In press). Among the more un- lemur species represented by wild-living popu- expected results of that study were that, at least lations outside Madagascar is Lemur mongoz seasonally, L. mongoz a) lives in "family" (=Lemurmongoz mongoz of most authors).' Such groups, containing an adult male and an adult populations are restricted to the Comoro Islands, female with or without immature offspring, and an archipelago of volcanic origin lying in the b) is nocturnal in its activity pattern. In both of northern part of the Mozambique Channel, be- these characteristics L. mongoz is unique among tween Madagascar and Africa (fig. 1). Of the four those members of its genus studied to date. islands comprising the archipelago, L. mongoz is From mid-November, 1974, to early January, found on two: Anjouan (424 km2) and Moheli 1975, a survey was undertaken of the L. mongoz (290 km2). Although the means of introduction populations of Moheli and Anjouan, during which of lemurs to the remains obscure, as attention was particularly directed toward group does its timing, this seems most likely to have composition and activity rhythm. This was done occurred as a result of human activities (Tatter- largely in order to test the possibility that noc- sall, In press). tumality and the formation of "family" groups The only previously published study of L. were seasonal or locale-specific in the Malagasy mongoz was carried out during July and August animals studied in 1973, and to augment the of 1973 at Ampijoroa, Madagascar, by the pres- rather narrow data-base of that study. ent author in collaboration with Dr. R. W. Suss- man (Tattersall and Sussman, 1975; Sussman and ACKNOWLEDGMENTS 'The species L. mongoz is generally considered to This study could not have been undertaken contain two subspecies: L. m. mongoz and L. m. coro- without the active co-operation of the Comorian natus. However, for a variety of reasons, which will be authorities. In particular, thanks are due the discussed elsewhere, it is clear that coronatus deserves following officials: in Moroni, M. Omar Tamou, separate specific status. Ministre du Developpement Rural; M. Ahmed 369 370 ANTHROPOLOGICAL PAPERS AMERICAN MUSEUM OF NATURAL HISTORY VOL. 53

FIG. 1. Location and disposition of the Comoro Islands in the Mozambique Channel.

Djabir, Directeur de l'Agriculture; M. Mouhtar Richard and Robert W. Sussman for reading and Rachide, Directeur de la Production, Eaux et commenting on it. Finally, my thanks above all Forets; Dr. Francis Debuissy, Veterinaire; and M. to Mle. Christine Lemery, who suffered through Robert Tapia, Conseiller Technique. In Anjouan: it all. Mm. Swabahaddine Ben Said Mohamed, Ibrahim Mohamed and Nadjib Ahmed, all of the Circon- PELAGE AND AGING CRITERIA scriptions Agricole or Forestiere, ; and the Prefet, M. Abdul Karim Said Omar; also In conditions of good visibility, discrimination M. Affane Mohamed, of . In Moh6li: M. between the sexes of Comorian L. mongoz posed le Prefet and M. le Chef de la Circonscription no problem because of a marked sexual di- Agricole, ; M. Mohamed Chararoumane, chromatism in the region of the head. Most use- Chef de l'Elevage; and M. Baco Mari, Surveillant fully, the cheeks and beard of the females (fig. 2) des Forets; also Mm. Halidi Combo and Ali are invariably white, whereas those of the males M'Lindri. Among private individuals, I especially (fig. 3) are a reddish brown. Additionally, the thank M. and Mme. J.-C. Favetto of Moroni; M. males possess gray faces and white muzzles, in and Mme. Soubreville, M. and Mme. Lavarec, and contrast to the black faces of the females, whose M. and Mme. Freville of Mutsamudu; and M. and muzzles usually possess a much less pronounced Mme. Latrille and M. and Mme. Ecrepont of whitish spot toward the tip. No evidence at all Fomboni. The study was made possible by a was found of the two color variants observed grant (no. 1305) from the National Geographic among males from the Madagascar mainland Society of Washington, D.C.; I am most grateful (Tattersall and Sussman, 1975). for the Society's generous support. The line The sexual distinction is less marked in terms drawings are the work of Ms. Marjorie Shepatin; I of body coloration, which is predominantly of a am grateful to Ms. Debra Anne Dobbs for her mid- to light-gray, usually with reddish brown help with the manuscript, and to Drs. Alison elements. It was found, however, particularly in 1 976 TATTERSALL: LEMUR MONGOZ 371 individuals from Anjouan, that whereas the vari- study to distinguish between male and female ably extensive reddish brown fur of the male juveniles and infants. tends to be restricted to the foreparts of the No direct information exists on the timing of animal, that of the females is generally concen- the breeding and birth seasons in Comorian, or, trated posteriorly, leaving the shoulders, fore- indeed, in Malagasy L. mongoz. Tattersail and limbs, neck, and top of the head a pure gray. Sussman (1975) found that the stages of develop- Resemblance to Malagasy L. mongoz is greatest ment of young L. mongoz in northwest Madagas- in animals from Moheli, where the mixture of car agreed with the assumption that the birth colors on the body is often somewhat more season of this form coincides with that of L. f: diffuse. Lemur mongoz from Anjouan, particu- fulvus in the same area; this would place the larly from the highlands, appears to be slightly birth season in mid-October. Extension of this larger than that from Moheli, and possesses a assumption to the Comoros suggests that the highly characteristic physiognomy. infant/juveniles observed in Moheli in mid- Infants and younger juveniles display the November were approximately four weeks of white cheeks and beard, and usually the dark age, and that those observed in Anjouan at the face, characteristic of the adult female (fig. 4). end of the study in early January were aged Only toward the very end of the survey were around 11 weeks. At the beginning of the study some reddish elements beginning to appear on the most precocious infants were beginning to the cheeks (but not yet on the throats) of some leave their mothers' backs and move around juvenile males. No attempt was made during the independently (transition from infant to juvenile

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.Ak.a FIG. 3. Adult male Lemur mongoz from Anjouan. status) for part of the time; again, if analogy with portion of occasions such as when the genital L. fulvus is permissible, this would suggest an age immaturity of males was observable. The cate- of four to five weeks since Sussman (In press) gory has not, therefore, been formally recognized recorded that at about four weeks of age the in the census tables presented here. young L. f. rufus begins to leave the mother's back. The transition to fully independent status THE ENVIRONMENT is a lengthy one, however; according to Sussman the infant L. f. rufus spends only 30 percent of Moheli, the smallest of the Comoros, is not its time away from the mother at 11-12 weeks, complex topographically. Roughly oval in shape, and becomes fully independent only at about 16 it consists of a central longitudinal ridge, 790 weeks. The margin of error on age-estimations meters in altitude at its highest point, which based on the degree of contact with the mother descends fairly sharply to the sea on either side. is thus large. Coastal plain is very narrow or nonexistent Subadult L. mongoz aged 14 to 16 months except in the low-lying eastern extremity of the have usually attained both adult size, or very island. Apart from certain limited areas of the close to it, and adult coloration, but are still central crest at altitudes in excess of 550 meters, sexually immature. Under prevailing observa- Moheli's vegetation consists entirely of secondary tional conditions they were thus difficult to dis- regrowth and plantations. These latter, while tinguish with certainty on more than a small pro- including ylang-ylang, bananas, manioc, and rice, 1 976 TATTERSALL: LEMUR MONGOZ 373 are primarily of coconut palms (Cocos nucifera), massifs rises to 1575 meters, and it is in this area, the dominant floral element of the island. Owing and in certain limited portions of the southern to the low density of human population peninsula, that the surviving rain forest typical of (approximately 40/km2), few regions of Moheli the Anjouan highlands (above about 700 meters) are entirely denuded of vegetation, and L. is concentrated. Such forest is disappearing at an mongoz is found throughout the island in relative ever increasing rate, but still supports L. mongoz abundance; the species is apparently at least as in what is probably the highest density any- successful here in secondary vegetation as in pri- where. In lower-lying areas of secondary vege- mary forest. tation, however, the abundance of the species is The much greater human population density lower than in comparable regions of Moheli, and of Anjouan (over 250/km2), in contrast, has led in the areas of most intensive ground-cover clear- to the removal of vegetal cover over large areas of ance L. mongoz is absent. the island. Anjouan may be visualized as three peninsulas arranged in a T-formation, striking north, west and south, and each with a more or METHODS less well-defined central ridge dropping sharply We followed two rules in censusing L. to the sea. In the central portion of the island, at mongoz: first, groups were not included in the the junction of the peninsulas, a complex of census results unless and until the observer was

FIG. 4. Juvenile female (left) and male Lemur mongoz from Moheli. Both are approximately six weeks old; note that both possess the white cheeks and beard characteristic of the adult female. 374 ANTHROPOLOGICAL PAPERS AMERICAN MUSEUM OF NATURAL HISTORY VOL. 53 certain of both the total number of animals groups censused are plotted in figure 5. Of a total active or resting together, and of the sex of each of 26 groups, eight consisted of one adult male noninfant or nonjuvenile. This normally involved and one adult female. Aside from groups A 10 following an active group for as long as was and A 16, which probably represent special cases practicable, and the careful searching of adjacent and which are discussed below, the remaining areas. In both Moheli and Anjouan the ratio of groups possessed either three (33%o) or four groups located to groups censused was more than (38%o) individuals. The third or fourth individual 2:1. Second, in order to minimize the possibility was an infant/juvenile in 50 percent of cases, and of double-counting, no area of forest was visited an animal of adult size in the same percentage. more than once unless no groups had been cen- Of these latter, 13 in total, six (46%) were males sused on the first visit, even where it was known and seven (54%o) were females. Mean group size that other groups were present. In the one case (excluding groups A 10 and A 16) was 3.1 indi- where an exception was made to this rule, the viduals. census results of the first visit were discarded. The only plausible interpretation of these Search time varied considerably according both to the terrain and to the density of lemurs TABLE 1 in the area being surveyed. Once a group was Localities and Compositions of Groups located, it was relatively rare that a marked flight of Lemur mongoz Censused on Anjouan respons9 was elicited. Resting groups often (Orthography of locality names is French, and moved siightly when the observers approached, follows where possible that of the 1959 which in fact aided exact counting; animals 1:50,000 map, Institut already active were variably easy to census Geographique National, Paris.) depending on their activity. Feeding animals were easiest to census; traveling ones the most Infant/ difficult, as their movement was relatively rapid Group Locality Male Female Juvenile and the terrain and vegetation often considerably A 1 Hayco 1 2 1 impeded observer movement. Few moving groups A2 Hayco 1 1 - were considered reliably censused if they had not A 3 Daouejou 1 1 1 been followed for at least 15 minutes; counts of A 4 Daouejou 1 1 1 resting groups were not recorded where the foliage A 5 Dzialandze 1 2 1 was dense except where movement, often A 6 Dzialandze 2 1 - awaited for a considerable period, confirmed that A 7 Dzialandze 1 2 all individuals had been counted. On the basis of A 8 M'trouni 2 1 1 a second study (Sussman and Tattersall, In press) A 9 M'trouni 1 2 it now seems reasonable to conclude that the L. A10 Jimilime - 2 1 2 1 mongoz groups studied in Madagascar are stable A 11 Pomoni A 12 Changani 1 1 - over in case of the data here time; the presented A 13 Hombo 1 1 1 on the there is more emphasis observer's A 14 Moujimouvia 1 1 - inferences as to the stability of the groupings A 15 Hagora Iamrogogo 2 1 1 seen. A 16 Bandankoa 2 2 1 light levels were measured using a Gossen A 17 Bandankoa 1 2 1 Luna Pro light meter with incident light screen. A 18 Chamninga 2 1 - Readings were taken at shoulder-level in areas A 19 Boueni 1 2 1 clear of vegetation with the sensor of the meter A 20 Dzialandze 2 1 1 oriented to the zenith. A 21 Daouejou 1 1 - A 22 Daouejou 1 1 A 23 Daouejou 1 1 GROUP COMPOSITION A 24 Daouejou 1 1 1 A 25 Hamnamgoum 2 1 1 The results of censusing of L. mongoz on A26 Maroheni 1 1 Anjouan are given in table 1, and the localities of 1976 TATTERSALL: LEMUR MONGOZ 375 figures is that, on Anjouan, L. mongoz forms where predation, even by man, is minimal. It "family" groups similar to those observed during may be significant in this connection that the the study in northwest Madagascar. Although observations reported here of a mean group size definite identification of subadults aged 14 to 16 of 3.1 individuals and of no infant mortality are months (the approximate equivalent of the "J2 identical with those reported by Pollock (1975) juveniles" of Tattersall and Sussman, 1975) was for Indri indri which, likewise, forms "family" possible in only a limited number of cases, the groups and was studied in an area where it was conclusion that the third adult-sized member of not hunted. any group was in fact subadult is strongly sup- The interpretation of Anjouan L. mongoz ported by the observation that the proportion of groups as "families" is yet further strengthened groups containing such members was identical by the fact that the sex ratio of "third," pre- with that of groups containing infant/juveniles. sumed subadult, animals is approximately 1:1. The implication of zero infant mortality may be This is what would be expected in a pair-bonding due to sampling error, but it may well be that species where, as here, there is no reason to sus- such mortality tends to be low in an environment pect any marked departure from a 1:1 sex ratio

FIG. 5. Outline of Anjouan, to show localities of groups censused. 376 ANTHROPOLOGICAL PAPERS AMERICAN MUSEUM OF NATURAL HISTORY VOL. 53 at birth, or differential viability between im- six groups in which one twin birth (all, on this mature males and females. reckoning, in 1973; none in 1974) would explain The two groups representing departures from the observed composition, only 73 percent con- the pattern generally observed in Anjouan may, form to a "family" structure. To explain all such as suggested earlier, best be regarded as special groups thus would, however, involve invoking an cases. Group A 10, consisting of two females, abnormally high rate of twinning. Moreover, one slightly smaller than the other, and living on among these 16 groups the ratio of presumed the outskirts of a large village, represents, accord- subadults (i.e., those that would have to be inter- ing to the testimony of villagers, a "family" preted as subadults to permit the inference of a group of which the male had recently been lost "family" group structure) to infant/juveniles is through human intervention. The composition of 1:1.45; this would represent an unexpectedly group A 16, consisting of two adult-sized males, large annual variation in either the birthrate or in two adult-sized females and a juvenile, may most the pattern of late juvenile mortality. plausibly be explained by twinning in 1973. This Of the remaining six groups, two contained would represent one birth in the 30 giving rise to three adult-sized males and two females; two the immature individuals censused on Anjouan, contained three adult-sized males, two adult- or 3.3 percent of births. Multiple births are not sized females, and one infant each. The fifth con- rare in the genus Lemur; for L. mongoz Hill tained four males, two females, and an infant; (1973) listed nine sets of twins out of 119 off- spring recorded, a rate of 8.2 percent twin births. TABLE 2 The assumption in this case of twinning would Localities and Composition of Groups of not, then, appear unreasonable. Lemur mongoz Censused on Moheli During the Madagascar study, counts of five (Orthography of locality names is French, and groups of L. mongoz yielded a mean of 2.6 indi- follows where possible that of the 1958 viduals per group: three of the five groups were 1:50,000 map, Institut adult pairs without offspring, one possessed four Geographique National, Paris.) including a nonadult in each age-cate- individuals, Infant/ gory, and the last contained two adults and a Group Locality Male Female Juvenile subadult. In 1974, three of these groups were recensused by Sussman (Sussman and Tattersall, M 1 M'ro Ouamaji 1 1 In press). The subadult female belonging to one M 2 Mibani 1 2 1 group in 1973 had disappeared, as expected, M 3 1 1 - whereas the juvenile male of 1973, now subadult, M4 Moihani 1 1 was still present. The other two groups re- M 5 Cariela 3 2 1 censused each contained a new juvenile Thus the M 6 Cariela 1 1 1 M 7 Comodjou 1 1 - census from which suggest that data Anjouan, M 8 Comodjou 4 2 1 on average, two each pair produces, surviving M 9 Djoiezi 3 2 - infants every three years, are in agreement with M 10 Domoni 2 2 1 the reproduction/early survival rates observed in M 11 Domoni 2 2 the protected reserve at Ampijoroa. It seems M 12 2 2 - likely, however, that survival rates in other, non- M 13 Singani 3 2 1 protected, areas of Madagascar are considerably M 14 Kangani 2 2 - lower. M 15 M'Babani 1 1 The Moheli census data (table 2), on the other M 16 M'Babani - 4 - M 17 Djoumadounia 3 2 - hand, provide a rather different picture. Of the M 18 Mahoudjani 1 2 1 22 groups censused, only 10 (45%) appear to M 19 Mdronisiri 2 2 - have been normally constituted "family" groups, M 20 Bandaharisiri 1 1 1 i.e., to have contained an adult pair, with or M 21 Darine 1 1 - without a maximum of two offspring, one in M 22 Combani 1 1 1 each age-category. Even if one adds to these the 1976 TATTERSALL: LEMUR MONGOZ 377 this was the largest agglomeration observed. The although local microclimatic variation (but least most oddly constituted of all, however, was of all on Moheli) may be substantial. In both group M 16, which contained four females. Such Moheli and Anjouan, November represents the a grouping is unlikely to have been other than tail end of the dry season; the generally rather ephemeral and, taken in conjunction with the sudden onset of the rains at the beginning of observation of a high proportion of pairs December has profound effects on the vege- together with groups of more complex com- tation. Not much rain fell during censusing on position, leads to the speculation that seasonal Moheli, but heavy rain began to fall immediately variations in group structure may occur, and that thereafter. If seasonal variation in group struc- censusing was undertaken during the transitional ture does occur, a season of climatic and pheno- period. logical change is a reasonable period in which to There are a few indications, albeit somewhat expect social change to take place. indefinite, in support of the hypothesis of Secondly, there is some observational evi- seasonal change, at least as far as Moheli is con- dence that might be viewed as supporting the cerned. First, if the observations reported here fission and/or coalescence of groups. Thus, when were, in fact, being made during a seasonal groups M 5 and M 6 were censused on November transition, the transition was occurring at a 23, the two were initially thought to constitute a logical time of year. There is a fairly sharp single unit: they were found resting very close to seasonal variation in rainfall in the Comoros, each other as dusk approached, without showing

FIG. 6. Outline of Moheli, to show localities of groups censused. 378 ANTHROPOLOGICAL PAPERS AMERICAN MUSEUM OF NATURAL HISTORY VOL. 53 any signs of unease. Finally, however, the however, that group structure shows geographical animals of group M 5 moved off by a route differences, i.e., that pair-bonding is typical of which took them closely by group M 6; the latter both Anjouan and Madagascar, but that in stayed where it was and showed no reaction Moheli the composition of social units is more whatever, leaving its resting place some 12 variable, possibly, but not certainly, on a minutes later, in almost the opposite direction. seasonal basis. What does clearly emerge is that Although in Madagascar home ranges of L. studies of entire annual cycles in different mongoz groups were observed to overlap exten- environments in all three areas will be needed sively, there and in Anjouan group encounters before we can confidently speak of group struc- involved a great deal of agitation. It thus appears ture in the species as a whole. unlikely that in either place totally discrete groups would rest in such close proximity yet ACTIVITY RHYTHM remain apparently indifferent to each other's presence. The most plausible explanation may be During the Madagascar study L. mongoz was that this incident (or nonincident) represented observed to be active solely at night, between either a large group in the process of fission, or a approximately 1800 and 0600 hrs. Ambient light stage in the coming-together of smaller units. intensities when activity commenced were in the But if this explanation is far from firm, deter- narrow range of 2.8 to 22.0 Lux, and when it mination of the polarity of such change is yet ceased, of 1.4 to 22.0 Lux. more so. Inference from L. mongoz elsewhere is In Madagascar, observations were made on hazardous in itself, and, beyond this, data from groups of L. mongoz that had become fully Anjouan and Madagascar, even if admissible, habituated to the observers. In the Comoros, on point in diametrically opposite directions. In the other hand, no intensive study preceded by Madagascar, pair-bonding was observed in L. an habituation period was made, and it was thus mongoz during the dry season; in Anjouan not not possible to be completely certain that only was it found at the beginning of the wet activity observed before nightfall was not season in environments apparently comparable stimulated by the arrival of the observers, usually with those surveyed in Moheli, but it was also in the company of local people. It seems proba- found to be characteristic of groups living in the ble that the lemurs were normally aware of the more or less permanently humid highlands of the presence of the observers well before we located interior. If such seasonal shifts do occur, it would them. Observational bias, therefore, is likely to seem more reasonable to expect the larger group- have been toward the early commencement of ings to characterize the wet season, the period of activity, although in general Comorian L. the greatest abundance of resources. On the basis mongoz is quite well habituated to human pres- of currently available data, however, the hypoth- ence and, since hunting is rare, not unduly dis- esis of seasonal change is far from certain; the turbed by it. possibility that we are observing geographical Surveying in the early morning was more diffi- variations due to factors such as habitat or popu- cult, and generally less successful, than in the lation density cannot be ignored. aftemoon and early evening. No L. mongoz were The overall sex ratios of adult-sized animals seen on Moheli to be active in the hours follow- were virtually identical between Moheli and ing sunrise, but sampling was unsatisfactory Anjouan: 37 males to 37 females (1:1) in the during this period, and attention will be paid former, and 32 males to 33 females (1:1.03) here only to observations made on that island (ignoring group A 10) in the latter. In Madagas- between noon and 2230. The official time of car, with a much smaller sample size, and includ- sunset during surveying on Moheli ranged be- ing the single JI juvenile, whose sex was ascer- tween 1807 hrs. and 1819, and on Anjouan tainable, the ratio was similar: six males to seven between 1822 and 1832. females (1:1.17). On Moheli, L. mongoz groups censused during It is, then, impossible at present to make any the hours of daylight were virtually always sleep- definite statement as to group composition, and ing, or, since they were generally aroused by the its variability, in L. mongoz. It seems most likely, approach of the observers, resting. During the 1976 TATTERSALL: LEMUR MONGOZ 379 late aftemoon juveniles were occasionally this observation was made was close to a well- observed moving around a little while the adults traveled path, it is possible that this early activity of their groups rested. On only two occasions was due to disturbance by earlier passersby; was L. mongoz observed to be active at light certainly, the later the hour at which sleeping L. levels higher than those which triggered activity mongoz groups are disturbed during daylight, the in the Malagasy animals. On November 24, at more likely they are immediately to commence Comodjou, group M 7, consisting of an adult sustained activity. male and female, was observed feeding at 1815, In the highland rain forest, however, at alti- at a light level of 115 Lux. On November 28, tudes above ca. 600-700 meters, the survey group M 15, another adult pair, was observed suggested a radically different activity pattern. traveling, although not rapidly, at 1746, when The difficulties of access to these areas precluded the light level was around 6300 Lux. On two the observation of L. mongoz later than about occasions L. mongoz groups were observed dur- 1930 hrs., but observations at other times indi- ing the late aftemoon in postures and cated that activity continued throughout the dispositions which did not suggest resting, hours of daylight and beyond. Of the 15 groups although the animals were not moving when censused in these highland areas between 0850 located. and 1930, only two were resting; a further two All L. mongoz observed in Moh6li after the may have been moving as a result of our official hour of sunset were active at the time of approach. All the others were moving, traveling censusing, generally either traveling or feeding. or feeding and, indeed, group encounters were No observations were made after 2230, and the observed on two occasions. One of these, activity invariably recorded between nightfall between groups A 8 and A 9, took place at and this time accords well with the observation M'trouni, at approximately 1230 on December of a prolonged activity period at Ampijoroa, 14, when the two groups, approaching from Madagascar, between approximately 1800 and opposite directions, surprised each other in the 2300. Further, observations on Moheli suggest same patch of vegetation. In form it was very that the pattern observed at Ampijoroa, where similar to the encounter witnessed at Ampijoroa initial activity consisted of a period of travel (Tattersall and Sussman, 1975), with the major away from the sleeping site followed by a long exception that vocalization, which at Ampijoroa feeding bout, held equally there. included "creaking door" vocalizations as well, Results from Moheli thus suggest that the pat- was limited to grunting, of which there was a tern of activity on this island does not differ sub- great deal. The other occurred toward dusk on stantially from that recorded in Madagascar. This December 30, at Daouejou, between groups A 23 corroborative observation allays the suspicion and A 24, and included the entire vocal reper- that the nocturnal activity of the animals toire heard at Ampijoroa. This was a more long- observed at Ampijoroa may have been related drawn-out confrontation than the earlier one, solely to the night-flowering of the kapok tree, lasting over a quarter of an hour as opposed to Ceiba pentandra, whose flowers provided their about two minutes, and commenced when the principal dietary resource during the period of light was already failing; it continued, in bouts, the study (Tattersall and Sussman, 1975). until the light level had sunk below 2.8 Lux, and The data from Anjouan, however, provide a it was followed by feeding sessions on the part of less clear-cut picture. In areas of low-lying both groups. secondary vegetation, which offer an environ- Ambient temperatures and light-levels are con- ment comparable with those surveyed on Moheli, siderably lower in the humid central highlands an essentially similar pattern was found to that than in the lower-lying coastal areas; at 1030 on observed on the smaller island. Only on one December 14, for instance, a light reading of occasion was a group (A 16) of L. mongoz seen only ca. 3400 Lux was recorded, this on an to be active earlier than 30 minutes before the average day for the time of year, and mist official hour of sunset; this was at Bandankoa at frequently descends over the rain forest. Since approximately 1715 on December 27, at a light diurnality was observed only under these level of ca. 1800 Lux. Since the locality at which environmental conditions, whereas in the drier, 380 ANTHROPOLOGICAL PAPERS AMERICAN MUSEUM OF NATURAL HISTORY VOL. 53 warmer, brighter and more seasonal lowland during the cold nights they experience. Again, as environments of both Anjouan and Moheli (and, in the case of group structure, longer-term for that matter, of Madagascar), a relatively strict studies in all three areas and in a variety of nocturnality was seen, variation in activity pat- environments will be necessary before the full tem appears to be environmentally linked. Possi- complexity and significance of activity patterns bly the diurnality of the lemurs of the Anjouan in L. mongoz can be appreciated. interior is due to limitations of thermoregulation

LITERATURE CITED Hill, Clyde A. Sussman, R. W., and I. Tattersall 1973. The frequency of multiple births in the [In press.] Cycles of activity, group composi- genus Lemur. Mammalia, vol. 37, no. 1, tion and diet of Lemur mongoz pp. 101-104. mongoz Linnaeus 1766 in Madagascar. Pollock, J. I. Folia Primat. 1975. Field observations on Indri indri: a pre- Tattersall, I. liminary report. In Tattersall, I., and R. [In press.] Note sur la distribution et sur la W. Sussman (eds.), Lemur Biology, situation actuelle des lemuriens des New York and London, Plenum, pp. Comores. Mammalia. 287-311. Tattersall, I., and R. W. Sussman Sussman, R. W. 1975. Observations on the ecology and be- [In press.] Socialization, social structure and havior of the mongoose lemur Lemur ecology of two sympatric species of mongoz mongoz Linnaeus (Primates, Lemur. In Chevalier-Skolnikoff, S., and Lemuriformes), at Ampijoroa, Madagas- F. Poirier (eds.), Primate Socialization. car. Anthrop. Papers Amer. Mus. Nat. Chicago, Aldine. Hist., vol. 52, pt. 4, pp. 195-216. ANTHROPOLOGICAL PAPERS OF THE AMERICAN MUSEUM OF NATURAL HISTORY

Volume 53 1976

PUBLISHED BY ORDER OF THE TRUSTEES NEW YORK: 1976 Edited by FLORENCE BRAUNER CONTENTS OF VOLUME 53 Part 1. Shanti Nagar: The Effects of Urbanization in a Village in North India. Stanley A. Freed and Ruth S. Freed. Pages

1-254, 19 figures, 114 tables. March 26, 1976 .... $13.85 Part 2. Notes on the Status of Lemur Macaco and Lemur Fulvus (Primates, Lemuriformes). Ian Tattersall. Pages 255-262, 2 figures. August 11, 1976...... $..65 Part 3. Prehistoric Pifion Ecotone Settlements of the Upper Reese River Valley, Central Nevada. David Hurst Thomas and Robert L. Bettinger. Pages 263-366, 65 figures, 7 tables. December 30, 1976 ...... $6.70 Part 4. Group Structure and Activity Rhythm in Lemur Mongoz on Anjouan and Moheli Islands, Comoro Archipelago. Ian Tattersall. Pages 367-380, 6 figures, 2 tables. December 30,

1976...... $1.15

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