The social contexts of some very low frequency calls of African elephants

Summary Several types of low frequency calls made by African elephants, Loxodonta africana, and the contexts in which they occurred are described. These calls had fundamental frequencies ranging from 14–35 Hz and sound pressure levels as high as 103±3dB (re 20 Pa) at 5 m from the source. Very low frequency sounds are subject to very little environmental attenuation, suggesting that sounds at the frequencies and sound pressure levels measured from elephants may be audible to conspecifics several km away. Long-term records on the behavior of elephants and on the contexts of specific call types suggest that elephants make use of infrasound in the spatial coordination of groups and as they search for mates.     

Spatial relationships and matrilineal kinship in African savanna elephant (Loxodonta africana) clans

African savanna elephants, Loxodonta africana, live in stable family groups consisting of adult females and their dependent offspring. During the dry season, clans consisting of several family groups typically share a common home range. We compared spatial relationships and mitochondrial DNA (mtDNA) haplotypes among 14 adult female elephants within 3 clans during the dry season in northern Zimbabwe. Spatial relationships were studied by radio-tracking. Home-range similarity was quantified by correlating the estimated utilization distributions of all pairs of elephants. Clans were identified by cluster analysis of the home-range similarity values. All three clans contained at least two of the five mtDNA haplotypes that were found, indicating that clan members are not necessarily matrilineally related. Within clans, home ranges of elephants with the same haplotype were not significantly more similar to each other than those of elephants with different haplotypes. Most elephants within each clan used their shared home ranges independently of each other: the distribution of distances between their positions at any given time did not differ from the distribution expected by chance. However, 8 out of the 26 within-clan pairs exhibited long-term coordination of space use by remaining within known hearing distance of each others low-frequency calls significantly more often than expected by chance. At least four of these coordinated pairs consisted of animals in different family groups. Elephants in three of the four different-family pairs whose movements were coordinated had different haplotypes. Further research is needed to determine the relationship between these coordinated movements and conventionally defined bond-group behavior.Electronic Supplementary Material Supplementary material is available in the online version of this article at .Communicated by C. Nunn     

Functional reference in an alarm signal given during nest defence: seet calls of yellow warblers denote brood-parasitic brown-headed cowbirds

Field observations and model-presentation experiments have shown that yellow warblers (Dendroica petechia) produce seet calls preferentially in response to brood-parasitic brown-headed cowbirds (Molothrus ater). In this study, we investigated whether seet calls are functionally referential alarm calls denoting cowbirds by determining whether female warblers responded appropriately to seet calls in the absence of a cowbird, whether alarm calling by warblers varied with response urgency, and how warblers in a population allopatric with cowbirds responded to cowbird and avian predator models and seet playbacks. As a control, we presented chip calls, which are elicited by nest predators as well as by non-threatening intruders, but are not strongly associated with cowbirds. Yellow warblers responded differently to playbacks of seet than chip calls. To seet playbacks, almost 60% of females gave seet calls and rushed to sit in their nests, responses typically elicited by cowbirds, whereas these responses were given infrequently in response to chip calls. Yellow warblers seet called equally in situations that simulated low, medium and high risk of parasitism, which suggests that these calls did not vary with response urgency. In a population allopatric with cowbirds, seet calls were rarely produced in response to cowbird or avian nest predator models and never to seet playbacks. These results suggest that seet calls are functionally referential signals denoting cowbirds and that cowbird parasitism was a strong selective pressure in the evolution of functional referentiality in the seet call of yellow warblers.Communicated by W.A. Searcy     

Molecular genetic and behavioral analysis of social organization in the Asian elephant (Elephas maximus)

We report on the genetic evaluation and behavioral study of social organization in the Asian elephant (Elephas maximus). Although Asian elephants and African elephants (Loxodonta africana) were previously thought to have similar social organizations, our results demonstrate a substantial difference in the complexity and structure of Asian elephant social groupings from that described for African savanna elephants. Photographic cataloging of individuals, radio telemetry, and behavioral observations in Ruhuna National Park, Sri Lanka, enabled us to assign associated females and young to four groups with overlapping ranges. Genetic sampling of individuals from the four groups in Ruhuna National Park and three other groups in surrounding areas, conducted through PCR amplification and sequencing of mitochondrial DNA from dung, supported the matriarchal nature of female groups and the lack of inter-group transfer of females. Behaviorally and genetically, the identified social groups were best described as ”family groups”. We did not find any evidence for the existence of social groups of higher complexity than family groups. Received: 25 March 2000 / Received in revised form: 28 March 2000 / Accepted: 1 April 2000     

Bat serenades—complex courtship songs of the sac-winged bat (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>)

Vocalisations of many songbirds, anurans, and insects are shaped by sexual selection. Males acoustically compete for territories, and females choose their mates by means of male courtship songs. In courtship, richness and complexity of elements are often favoured characters. Only a few examples of complex songs are known in mammals. Males of the harem-polygynous sac-winged bat (Saccopteryx bilineata, Emballonuridae) have an uncommonly complex vocal repertoire, and different song types of males are used in the context of territorial defence and in courting females. We classified the daytime vocalisations of 16 male S. bilineata from a colony in Costa Rica, both on the basis of their acoustical properties and the social context in which they occurred. Seven vocalisation types were differentiated: echolocation pulses, barks, chatter, whistles, screeches, territorial songs and courtship songs. Territorial songs were short, rather stereotyped and not obviously directed towards a certain conspecific. They appear to be of importance in male competition for harem territories, in which females roost during the day. Courtship songs were exclusively observed when males displayed towards a female; they were long and complex, and consisted of highly variable elements (calls). We classified the calls in courtship songs of six males into call types, based on acoustical properties, mainly spectral purity and duration. Four call types are described in detail: trills, noise-bursts, short tonal calls, and quasi constant frequency calls. Twelve parameter values were extracted from the most common call type, the trill. Discriminant function analysis of trills showed that different males had different repertoires. This could allow females to use trill parameters for recognition of individual males and thus for mate choice.Communicated by G. Wilkinson     

Ground gleaning in horseshoe bats: comparative evidence from<Emphasis Type="Italic"> Rhinolophus blasii</Emphasis>,<Emphasis Type="Italic"> R. euryale</Emphasis> and<Emphasis Type="Italic"> R. mehelyi</Emphasis>

The 71 species of horseshoe bat (genus Rhinolophus) use echolocation calls with long constant-frequency (CF) components to detect and localize fluttering insects which they seize in aerial captures or glean from foliage. Here we describe ground-gleaning as an additional prey-capture strategy for horseshoe bats. This study presents the first record and experimental evidence for ground-gleaning in the little-studied Blasius horseshoe bat (Rhinolophus blasii). The gleaning bouts in a flight tent included landing, quadrupedal walking and take-off from the ground. The bats emitted echolocation calls continuously during all phases of prey capture. Both spontaneously and in a choice experiment, all six individuals attacked only fluttering insects and never motionless prey. These data suggest that R. blasii performs ground-gleaning largely by relying on the same prey-detection strategy and echolocation behaviour that it and other horseshoe bats use for aerial hawking.We also studied the Mediterranean horseshoe bat (R. euryale) in the flight tent. All four individuals never gleaned prey from the ground, though they appeared to be well able to detect fluttering moths on the ground. It is not known yet whether ground-gleaning plays a role in Mehelys horseshoe bat (R. mehelyi). In a performance test, we measured the ability of these three European species of middle-sized horseshoe bats (R. euryale, R. mehelyi and R. blasii) to take-off from the ground. All were able to take flight even in a confined space; i.e. the willingness to ground-glean in R. blasii is not related to a superior take-off performance. In contrast to ground-gleaning bats of other phylogenetic lineages, R. blasii appears not to be a specialist, but rather shows a remarkable behavioural flexibility in prey-capture strategies and abilities. We suggest that the key innovation of CF echolocation paired with behavioural flexibility in foraging strategies might explain the evolutionary success of Rhinolophus as the second largest genus of bat.Communicated by T. Czeschlik     

Foraging behavior and echolocation of wild horseshoe bats Rhinolophus ferrumequinum and R. hipposideros (Chiroptera,Rhinolophidae)

Summary 1. Echolocation and foraging behavior of the horseshoe bats Rhinolophus ferrumequinum and R. hipposideros feeding under natural conditions are described. 2. The calls of both species consisted predominantly of a long CF segment, often initiated and terminated by brief FM sweeps of substantial bandwidth. 3. R. hipposideros typically flew close to vegetation, and fed by aerial hawking, gleaning and by pouncing on prey close to the ground. R. hipposideros called with a CF segment close to 112 kHz which is the second harmonic of the vocalization; its calls included low intensity primary harmonics, and had prominent initial and terminal FM sweeps of considerable bandwidth. When searching for prey on the wing it had longer interpulse intervals than R. ferrumequinum, but emitted shorter pulses at a higher repetition rate; overall it had a similar duty cycle to R. ferrumequinum. 4. R. ferrumequinum, calling with a CF segment close to 83 kHz, also used harmonics other than the dominant secondary in its calls, and modified its echolocation according to ecological conditions. This species showed certain parallels with R. rouxi of Asia. It was observed feeding by aerial hawking and by flycatching. When scanning for prey from a perch (perch hunting), calls were of shorter duration, and interpulse intervals were on average longer, than when bats were flying. Mean duty cycle was longer in flight, and the bandwidths and frequency sweep rates of the FM segments in the calls increased in comparison with perched bats. 5. FM information may facilitate determination of target range and the location and nature of obstacles; it may also be involved in the interpretation of echoes and the detection of moving targets among clutter. The rising FM sweep initiating the call in both species when flying (and to a lesser extent perch hunting) in the wild must have a significant adaptive role, and should be considered an essential component of the call; rhinolophids should be termed FM/CF/FM bats.Abbreviations CF constant frequency - FM frequency modulated - FM₁ initial rising frequency sweep - FM₂ terminal falling frequency sweep - PRR pulse repetition rate - SD standard deviation - SNR signal-to-noise ratio     

The hearing of an avian predator and its avian prey

Summary Auditory tuning curves of a small songbird, the great tit (Parus major), and of its principal avian predator, the European sparrowhawk (Accipiter nisus), were determined by an operant positive reinforcement conditioning procedure, using the method of constant stimuli. Thresholds were measured by the criterion of a 50% correct response and a d of 1.5 for intra- and interspecific comparison, respectively. The best frequency of both species was 2 kHz, the hawk being 6.5 dB SPL more sensitive than the tit. Although the high-frequency cutoff was very similar in both species, at 8 kHz the great tit was about 30 dB more sensitive than the sparrowhawk. The hearing abilities of the prey and its predator are discussed with reference to the acoustic alarm communication of great tits confronted with sparrowhawks. Two alarm calls lie in the frequency range of the best hearing of both the hawk and the tits: the mobbing call and a call given in response to a nearby hawk when fleeing from it. In contrast, the seeet call, an alarm call given mainly in response to distant flying sparrowhawks, can only be heard well by the tit. The implications of these results for hypotheses concerning the evolution of alarm calls in small songbirds are discussed.     

Long‐Term Impacts of Poaching on Relatedness,Stress Physiology,and Reproductive Output of Adult Female African Elephants

Abstract: Widespread poaching prior to the 1989 ivory ban greatly altered the demographic structure of matrilineal African elephant (Loxodonta africana) family groups in many populations by decreasing the number of old, adult females. We assessed the long‐term impacts of poaching by investigating genetic, physiological, and reproductive correlates of a disturbed social structure resulting from heavy poaching of an African elephant population in Mikumi National Park, Tanzania, prior to 1989. We examined fecal glucocorticoid levels and reproductive output among 218 adult female elephants from 109 groups differing in size, age structure, and average genetic relatedness over 25 months from 2003 to 2005. The distribution in group size has changed little since 1989, but the number of families with tusked old matriarchs has increased by 14.2%. Females from groups that lacked an old matriarch, first‐order adult relatives, and strong social bonds had significantly higher fecal glucocorticoid values than those from groups with these features (all females R²= 0.31; females in multiadult groups R²= 0.46). Females that frequented isolated areas with historically high poaching risk had higher fecal glucocorticoid values than those in low poaching risk areas. Females with weak bonds and low group relatedness had significantly lower reproductive output (R²[U]=0.21). Females from disrupted groups, defined as having observed average group relatedness 1 SD below the expected mean for a simulated unpoached family, had significantly lower reproductive output than females from intact groups, despite many being in their reproductive prime. These results suggest that long‐term negative impacts from poaching of old, related matriarchs have persisted among adult female elephants 1.5 decades after the 1989 ivory ban was implemented.     

Evolutionary genetics of the Mytilus edulis complex in the North Atlantic region

Genetic relationships among Mytilus populations throughout the North Atlantic region, including the Mediterranean and the Baltic Sea, were studied using enzyme electrophoresis. Three distinct groups of populations, each of a remarkably wide distribution, can be recognised on the basis of their multilocus allelic composition: (1) M. galloprovincialis L. of the Mediterranean and western Europe; (2) a genetically distinct form of M. edulis Lmk. from both the Baltic Sea and some localities in the Canadian Maritime Provinces (here provisionally termed the trossulus type mussel); and (3) the traditional Atlantic M. edulis populations of northwestern European coasts and most of eastern North America. These groups are regarded as representing three relatively old evolutionary lineages, which all deserve separate and equal systematic status. The main part of the differentiation at most of the loci studied is accounted for by this major systematic pattern, but considerable geographical differentiation within each of the three principal groups was also detected. At single loci, different electromorphs were found to prevail in disjunct populations of M. galloprovincialis (Mediterranean/Britain) and of the trossulus-type mussel (Baltic/eastern Canada). Within the Atlantic M. edulis, a major part of the differentiation is transoceanic. At one locus (Ap), geographic differentiation appeared to be relatively independent of the systematic boundaries; the possible role of interlineage hybridisation in contact areas in regulating the pattern of geographical variation is discussed.     

β-glucanase activity in the seastar Pisaster ochraceus

Specimens of Pisaster ochraceus, an intertidal carnivorous seastar, were collected in northern Puget Sound at Post Point, near Bellingham, Washington, USA, in November 1993, and used in experimentation through September 1994. Ammonium sulfate fractions (60% precipitates) from excised pyloric caeca (digestive glands) possessed -glucanase activity directed toward such -linked substrates as carboxymethylcellulose (CMC), partially-digested cellulose (cellodextrins) and the synthetic substrate p-nitrophenyl--D-glucoside (PNDG). pH-activity and pH-stability maxima were at pH 6 and 5, respectively. No hydrolytic activity was detectable on native cellulose, native chitin or on cationic-modified celluloses such as DEAE-cellulose. Gel filtration (Sephadex G-100) of this 60% precipitate followed by DEAE-Sephadex column chromatography revealed heterogeneity of -glucanase activity, with an indication of up to 11 distinct fractions with variable activity directed toward the substrates indicated above (CMC, cellodextrins and PNDG). These fractions displayed differing ratios of hydrolytic activity on these substrates. None of the isolated fractions displayed activity toward either native cellulose or native chitin. Gut extracts from the edible mussel (Mytilus edulis), a common prey species of P. ochraceus, were shown to possess partially digested cellulose (cellodextrins). These cellodextrins were further degraded by seastar -glucanases in vitro. The digestion of the gut contents of ingested prey provides a possible adaptive explanation for the presence of -glucanase activity in a carnivore.     

More accurate chronological age determination of crustaceans from field situations using the physiological age marker,lipofuscin

A captive population of Australian red-claw crayfish, Cherax quadricarinatus, of known age was used for the study. Lipofuscin concentration in the left olfactory lobe cell mass of the brain was measured using image analysis of fluorescence micrographs. Age predictions based on lipofuscin concentration were more accurate and had narrower 95% confidence limits than those derived from the more conventional body size and weight predictors for all age groups tested (180, 300, 420, 780 d). Overall, use of lipofuscin concentration produced a significantly lower (p0.001) mean age-prediction error (16.65%) than use of carapace length or body weight (32.45 and 32.3%, respectively). Carapace length was of little value for prediction of the age of older individuals. Mathematical models defining the relationships between temperature, age and lipofuscin accumulation were derived from laboratory-reared individuals. These models did not describe adequately the course of lipofuscin accumulation in crayfish from the field over the whole lifespan. Field data from three known year-classes were used to generate simulated size-frequency and lipofuscin concentration-frequency histograms. Year-classes in the lipofuscin concentration frequency-histogram were much more easily distinguished than in the size-frequency histogram. Under field conditions, lipofuscin concentration was a better predictor of chronological age than conventional morphometric measures.     

The role of alarm responses in the formation of mixed-species flocks of heathland birds

Summary Birds of ten species with similar short flight distances from a human intruder associated with stonechats (Saxicola torquata), which have large flight and alarm call distances. The attendants probably took advantage of the domain of safety around the stonechat, produced by the difference in flight distance. Stonechats have no reciprocal advantage, and were not seen to initiate flocks.Flocks were formed with both calling and silent stonechats. Benefits in the latter case probably include early warning of danger, but there may also have been a reduction in vigilance by attendants.The relative flocking activity of attendant species (number of associations observed; percentage of incidents with silent stonechats; rate of warning calls by stonechats joined) was related to the species' average flight distance from an intruder (Fig. 1). Possible explanations are consistent with a correspondence between flocking tendency and the degree of benefit obtained.     

The effect of perceived intruder proximity and resident body size on the aggressive responses of male green frogs, Rana clamitans (Anura: Ranidae)

We examined variation in aggressive responses within and among individuals in the green frog, Rana clamitans. We tested the hypothesis that resident males respond in a graded fashion to changes in perceived intruder proximity (stimulus intensity). We also investigated how response level varied with responder body size. We found that green frogs differentially alter aggressive responses (increasing movement and calling rates while decreasing the duration and dominant frequency of their calls) with an increase in stimulus intensity. Body size did not appear to be a significant influence on most response levels. We suggest that aggressive responses are graded, and advertisement and aggressive calls represent opposite ends of a continuum. Green frogs do not exhibit clearly defined thresholds for responses like those reported for some other frogs. By using graded responses rather than discrete all-or-nothing responses, males may reduce the potential costs of aggressive encounters.     

Vocal mediation of foraging competition in the cooperatively breeding green woodhoopoe (<Emphasis Type="Italic">Phoeniculus purpureus</Emphasis>)

Many group-living species produce frequent vocalisations when foraging, but the function of these food-associated calls is often difficult to divine. I investigated the kek call of the cooperatively breeding green woodhoopoe (Phoeniculus purpureus), a species in which individuals have preferred foraging techniques dependent on their bill size. Individuals called at a greater rate (1) in foraging compared to non-foraging situations, and (2) in groups containing potential foraging competitors (i.e. individuals that foraged using the same preferred techniques). I therefore asked whether the kek call is used to recruit conspecific foragers or whether it acts as a vocal signal of foraging niche and mediates foraging competition. Foragers that were vocalising were no more likely to be approached than those that were silent, and individuals gained no foraging advantage from the close proximity of another group member. Thus, keks are unlikely to be used to recruit conspecifics. Instead, they appear to regulate spacing between potential foraging competitors. Although an individual forager was equally likely to be closely approached by all other group members, it increased its calling rate only in response to potential foraging competitors. This increase in calling rate resulted in the approaching individual moving away, thus maintaining some separation between individuals that forage in the same way. Maintenance of such spacing is important because the success rate of an individual decreased when a foraging competitor was close by.Communicated by M. Leonard     

Functional nonredundancy of elephants in a disturbed tropical forest

Conservation efforts are often motivated by the threat of global extinction. Yet if conservationists had more information suggesting that extirpation of individual species could lead to undesirable ecological effects, they might more frequently attempt to protect or restore such species across their ranges even if they were not globally endangered. Scientists have seldom measured or quantitatively predicted the functional consequences of species loss, even for large, extinction‐prone species that theory suggests should be functionally unique. We measured the contribution of Asian elephants (Elephas maximus) to the dispersal of 3 large‐fruited species in a disturbed tropical moist forest and predicted the extent to which alternative dispersers could compensate for elephants in their absence. We created an empirical probability model with data on frugivory and seed dispersal from Buxa Tiger Reserve, India. These data were used to estimate the proportion of seeds consumed by elephants and other frugivores that survive handling and density‐dependent processes (Janzen‐Connell effects and conspecific intradung competition) and germinate. Without compensation, the number of seeds dispersed and surviving density‐dependent effects decreased 26% (Artocarpus chaplasha), 42% (Careya arborea), and 72% (Dillenia indica) when elephants were absent from the ecosystem. Compensatory fruit removal by other animals substantially ameliorated these losses. For instance, reductions in successful dispersal of D. indica were as low as 23% when gaur (Bos gaurus) persisted, but median dispersal distance still declined from 30% (C. arborea) to 90% (A. chaplasha) without elephants. Our results support the theory that the largest animal species in an ecosystem have nonredundant ecological functionality and that their extirpation is likely to lead to the deterioration of ecosystem processes such as seed dispersal. This effect is likely accentuated by the overall defaunation of many tropical systems.     

Gene flow among giant clam (Tridacna gigas) populations in Pacific does not parallel ocean circulation

Tridacna gigas (L.) populations were sampled from 13 locations throughout the west Pacific in 1990–1991 and screened for allozyme variation at seven polymorphic loci. Significant genetic differences were found between the few large stocks of T. gigas remaining in the world. Cluster analysis identified two major regional groups of populations: a Central Pacific group consisting of Kiribati and the Marshall Islands, and a West Pacific group comprising all other populations. Within the latter group, the Great Barrier Reef populations were significantly differentiated from those in the Solomon Islands. The main routes of gene flow did not run parallel with known major surface currents as might be expected, but crossed them. Gene exchange was limited east-west between the Central and West Pacific groups and the GBR and the Solomon Islands. The lack of correspondence between the major surface currents and the routes of gene flow suggests that the genetic structure of T. gigas reflects historical patterns of migration that no longer occur, rather than presentday dispersal. These findings raise fundamental questions concerning the origin and maintenance of genetic diversity in Pacific faunas, and have important implications for the mariculture and conservation of T. gigas.     

Roles of interplant movement,acoustic communication,and phototaxis in mate-location behavior of the leafhopper Graminella nigrifrons

Summary Male Graminella nigrifrons leafhoppers (Cicadellidae: Homoptera) employ a call-fly strategy to find virgin females on oat host plants. Males observed in isolation during daylight hours exhibit a high rate of interplant movement, calling from the lower canopy on each plant visited. Virgin and mated females exhibit little interplant movement. They differ from one another in that virgin females perch on the upper half of plants, whereas mated females perch on the lower half of plants. The positioning of females in the plant canopy is influenced by light. Unlike mated females, virgin females respond to male calls by emitting their own acoustic signals. When virgin females are present on plants visited by males, interplant movement of males ceases, and a localized-upward search of the female bearing plant ensues. Male search is influenced by light. Regardless of whether virgin females were confined to the upper or lower portion of plants, direction of male search was towards a light source used to illuminate above or below the plant canopy. These findings suggest that interplant movement by males and sedentary behavior by females prior to mate recognition and their use of acoustic and phototactic sensory modalities after mate recognition represent previously unrecognized adaptations to problems associated with the use of vibrational signals on plants. Offprint requests to: R.E. Hunt at the current address     

Gonadal development in the coral reef damselfish<Emphasis Type="Italic"> Dascyllus flavicaudus</Emphasis> from Moorea,French Polynesia

The present study investigated the sexual pattern of the yellow-tailed dascyllus, Dascyllus flavicaudus (Randall and Allen), through histological examination of the process of gonadal differentiation and maturation and through male-removal experiments in the laboratory. The study was conducted in Moorea, French Polynesia (17°32S; 149°50W), from 31 December 1994 to 11 March 1995. For gonadal histology, two populations were sampled, one consisting of large aggregations (n=54) and the other consisting of small- to medium-sized groups around isolated corals (n=55). An additional small sample (n=21) was also collected from a population that consisted of small groups around isolated corals. After an initially undifferentiated state, gonads of D. flavicaudus developed an ovarian lumen, followed by oocytes in the primary growth stage. From this ovarian state or from more developed ovaries with cortical-alveolus stage oocytes, some gonads developed into testes through degeneration of oocytes and development of spermatogenic tissue. In all three populations, spermatogenic tissue developed only in gonads that contained pre-vitellogenic oocytes (termed as mixed stage 1–2 gonads). The two main populations did not differ in their expression of sex change despite differences in their social organization. In both populations, size of individuals with mixed stage 1–2 gonads overlapped mainly with the size range of immature females, which suggests that functional female-to-male sex change was rare. The hypothesis that D. flavicaudus is primarily gonochoristic is further supported by removal experiments in laboratory groups, in which removal of a dominant male(s) failed to induce sex change in any of the high-ranking females. Yet, in all three field populations, some fish with mixed stage 1–2 gonads were found within the size ranges of mature females, which indicates possible occurrences of functional sex change. These conflicting results indicate that it may be premature to draw any definitive conclusions about the sexual pattern of D. flavicaudus.Communicated by T. Ikeda, Hakodate     

The neglected scent: on the marking function of urine in Lemur catta

In ring-tailed lemurs (Lemur catta) urine deposition can be combined with two different tail configurations: (i) tail held up in an evident display (urinate tail-up, UT-up); (ii) tail slightly raised to avoid its impregnation with urine (urinate tail-down, UT-down). We used both signaller- and receiver-based approaches to search for functional differences between these two kinds of urine deposition. We predicted that UT-up might be a complex signal combining olfactory and visual cues. We carried out observations and scent tests on four captive groups of ring-tailed lemurs. Group members sniffed/licked UT-up scents more frequently than UT-down ones. Moreover, UT-up showed peak levels during the mating season whereas UT-down did not. These findings suggest that urine can play a role in intra-group reproductive communication. Lemurs more frequently performed UT-up in a few drops and UT-down in streams. Recognition experiments clearly showed that individuals can discriminate between urine of their own group and urine from a foreign group (a necessary prerequisite for the use of urine in inter-group communication). The possible function of UT-up in inter-troop communication was supported by the higher frequency of this pattern along a fence separating two of the study groups. Moreover, in the presence of a dummy, the frequency of UT-up increased significantly. In conclusion, UT-up is a complex signal with multiple characteristics. By using different sensory channels, UT-up provides different types of information (location and signaller quality) and contains multiple messages directed both at group-members and neighbouring groups....to conceal this letter, the Minister had resorted to the comprehensive and sagacious expedient of not attempting to conceal it at all (from the Purloined letter E. A. Poe, 1845)