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     

A Population Viability Analysis for African Elephant (Loxodonta africana): How Big Should Reserves Be?

We present an age-structured, density-dependent model of elephant population dynamics in a fluctuating environment, drawing primarily upon the life history parameters obtained from studies in semi-arid land at Tsavo National Park, Kenya. Density regulation occurs by changes in the age of first reproduction and calving interval. We model environmental stochasticity with drought events affecting sex- and age-specific survivorships. Results indicate a maximum population growth rate of 3% per year and an equilibrium elephant density of 3.1/mile². Analysis of the demographic results and their sensitivity to changes in juvenile survivorship and drought frequencies, supported by genetic considerations, suggests that in semi-arid regions a minimum reserve size of 1000 mile² is necessary to attain a 99% probability of population persistence for 1000 years. The effect of age-independent culling on population viability is also analyzed.     

Male dominance,female choice and male copulatory behavior in two species of voles (Microtus ochrogaster and Microtus montanus)

Summary Female preferences for dominant males in prairie and montane voles were analyzed in two different test situations. In the first, prairie vole females preferentially spent time in proximity to dominant versus subordinate males which were housed behind a wire mesh screen. In a two-male tether test prairie voles females both preferentially associated and mated with dominant males. Montane voles, on the other hand, showed no preference in either test situation. The baseline copulatory behavior of naive montane vole males which became dominant differed significantly from those which became subordinate; no such differences were evident in the baseline copulatory behavior of naive praire voles. One hour following dominance testing, there was no difference evident between dominant and subordinate males of either species but there was a general facilitation of male copulatory behavior in both species, the effect being of greater magnitude in montane voles.     

Maternal quality and differences in milk production and composition for male and female Iberian red deer calves (Cervus elaphus hispanicus)

Several theories predict a sex-biased investment either through unbalanced sex ratios in offspring or through differences in provisioning. According to them, one would expect an optimisation in indirect fitness, or else a compensation for increased mortality of one sex. In addition, biases in provisioning may also arise as a consequence of weight-dependent non-adaptive nutrient demands by offspring. This study examines milk provisioning and sex biases in offspring sex ratio together with maternal quality variables. Mothers of higher quality (weight and age) showed greater milk provisioning ability (in terms of production) resulting in greater calf weight gain. Mothers of sons produced greater yields of milk, milk protein, fat and lactose than mothers of daughters, and increased percentage of protein after controlling for higher male birth weight. In contrast, mothers of males did not differ from mothers of females in age or any body weight variables related to maternal quality. These results suggest that differences in milk production and composition for sons and daughters are rather a mechanism to optimise indirect fitness than a mechanism to compensate for increased mortality in male calves, or a consequence of greater weight-dependent nutrient demands by heavier male calves. Results also suggest that biases in milk provisioning may occur without biases in offspring sex ratio, and furthermore, in contrast to the prediction that biases should be relative to the mean investment of the population, that milk provisioning biases might not be relative.Communicated by F. Trillmich     

Effects of movement and eating on chemosensory tongue-flicking and on labial-licking in the leopard gecko (Eublepharis macularius)

Summary Two forms of lingual protrusion, tongueflicking and labial-licking, were differentially affected by combinations of movement and eating conditions in a eublepharid gecko (Eublepharis macularius). Tongue-flicking, in which the tongue contacts substrates beyond the lizard's body, occurred at increased rates during locomotion and during locomotion was significantly more frequent after eating than in a baseline condition. Labial-licking, in which a protruded portion of the tongue touches the labial, mental or rostral scales that surround the mouth, increased after eating. Unlike tongue-flick rates, by far the highest labial-lick rates were observed in stationary lizards after eating. The elevated tongue-flicking rates during movement after eating may be a manifestation of a postingestive chemosensory search for prey. In addition to grooming, several possible chemosensory functions of labial-licking are discussed, including gustatory sampling, sampling prey chemicals on the labials for transfer to the vomeronasal system, and redistribution of chemicals on the tongue to enhance transfer. It is suggested that labial-licking might help motionless lizards maintain vigilance for visual prey stimuli associated with the specific chemical prey cues. Another possible explanation for the increased labial-lick rate while motionless after eating is that prey chemicals induce tongue-flicking, but that the distance protruded is lessened and the tongue does not contact environmental substrates. Tongue-flicking while stationary is unlikely to lead to detection of additional prey and might incur detection by the lizard's predators or prey.     

Feeding by larvae of two different developmental modes in Streblospio benedicti (Polychaeta: Spionidae)

Females of the spionid polychaete Streblospio benedicti (Webster) produce either small eggs (60–70 μm diameter) and planktotrophic larvae, or large eggs (100–200 μm) and lecithotrophic larvae that reportedly do not feed. This intraspecific polymorphism, a form of poecilogony, is potentially useful in studies of larval ecology and evolution, but necessary data on larval form and function are lacking. This study describes the morphology and nutritional biology of larvae obtained from Atlantic (South Carolina) and Pacific (California and Washington) populations from 2003 to 2005. The two types of larvae produced by Atlantic S. benedicti differed greatly in length (229±22 μm SD for planktotrophs vs. 638±40 μm for lecithotrophs) and chaetiger number (2–5 vs. 10–11) at release from the female’s brood pouch. Planktotrophic larvae bore long provisional chaetae on their first chaetiger; provisional chaetae were absent in lecithotrophic larvae. Larvae from Pacific populations were all of the lecithotrophic form, and were similar to their Atlantic counterparts in all respects. High-speed video microscopy revealed that both types of larvae used opposed bands of cilia to capture suspended particles and transport them to the mouth, where they were often ingested. Lecithotrophic larvae reared with suspended phytoplankton (Rhodomonas sp., 10⁴ cells ml⁻¹) for 2 days grew significantly faster than sibling larvae reared without added food, indicating that these larvae can digest and assimilate ingested food. Larvae of S. benedicti that develop from large eggs are thus facultative planktotrophs instead of obligately non-feeding lecithotrophs, a result that affects the interpretation of comparative studies of the ecology and evolution of larvae in S. benedicti and certain other marine invertebrates.     

Male takeover,infanticide, and female countertactics in white-headed leaf monkeys (<Emphasis Type=Italic>Trachypithecus leucocephalus</Emphasis>)

Infanticide by males is common in mammalian species such as primates in which lactation lasts much longer than gestation. It frequently occurs in one-male groups following male takeovers and is likely a male reproductive strategy. Reported female countertactics include abrupt weaning of infants, dispersal, or paternity confusion. Here, we estimated costs of female countertactics in terms of weaning ages and interbirth intervals. We observed a population of white-headed leaf monkeys (Trachypithecus leucocephalus) in Nongguan Nature Reserve, China (1995–2006) mainly composed of one-male groups. Takeovers (N = 11) coincided with the peak conception period. Detailed data are presented for five takeovers (34 females, 29 infants, and 47 group-years) leading to six infant disappearances (42.9% of infant mortality). All presumed infanticides were in accordance with the sexual selection hypothesis. Following a takeover, females without infants or with old infants stayed with the new males, incurring no or low costs (via abrupt weaning). Females with young infants dispersing with the old males also experienced low costs. High costs (due to infant loss) were incurred by pregnant females and those with young infants who stayed with the new males indicating that paternity was not confused. Costs in terms of long interbirth intervals were also high for females leaving with the old males to later join the new males, despite infant survival. Female countertactics reflected female philopatry mediated by infant age. Presumably due to the seasonal timing of takeovers, most countertactics seemed successful given that 32.3% of females apparently incurred no costs and 41.2% incurred only low costs.     

Choice and utilization of oviposition sites by female Calopteryx maculata (Odonata: Calopterygidae)

Summary Females of the damselfly Calopteryx maculata (de Beauvois) initially choose the larger of a pair of adjacent oviposition sites, about 70% of the time (Table 1), or whichever of two equal sized sites had other ovipositing females on it (about 88% of the time-Table 2). These criteria for initial choice between a pair of sites also interact. Incoming females generally (57 to 74% of the time-Table 3) joined others on the small site rather than ovipositing alone at the adjacent, bigger site. When pairs of large and small sites were replicated across eight locations, there were nonsignificant trends towards greater utilization (eggs laid) of the larger of a pair of oviposition sites within locations (Table 4). The lack of agreement between initial choice and utilization shows that other factors besides size are important in the choice and use of oviposition sites. These include disturbance by males, the presence of other females and choice criteria that can only be assessed during oviposition. When all sites at the eight locations were equal in size, there was considerable day to day and location to location variation in eggs laid (Fig. 1). Viewed over periods of several days, some sites are obviously less attractive than others in terms of cumulative numbers of eggs laid at them. When the amount of vegetation was varied among locations, those with the bigger oviposition sites were used more often, somtimes significantly so, but there were also significant reversals (small sites used more often) (Table 5, Fig. 1). Thus, there is no simple effect of size on the utilization of oviposition sites by Calopteryx maculata females, despite a clear tendency for females to make initial choices based on this criterion. The considerable among and within location variation in number of eggs laid may reflect additional choice criteria or the interaction of size, the presence of other females, disturbance, and location.     

Tail size and female choice in the guppy (Poecilia reticulata)

Summary Under laboratory conditions, female guppies demonstrate a clear preference for males with larger tails, and this preference translates into enhanced reproductive fitness for these males. Females also prefer males with higher display rates, a behavior which appears to be linked to tail size, but which can be experimentally disassociated. This appears to be a case of female-choice sexual selection.     

Male mating success and paternal care in Nicrophorus vespilloides (Coleoptera: Silphidae)

Summary If no female is present, male burying beetles Nicrophorus vespilloides (Coleoptera: Silphidae) co-operate in the burial of a corpse. Once a female has arrived, the males fight with one another. The defeated male stays near the corpse and to copulate with the female. Laboratory experiments using sterilised males showed that the defeated male was able sometimes to father some of the offspring raised on the corpse. Male N. vespilloides almost always participate in defence and feeding of the brood. This is not affected by the size of the male. Males quickly leave or are driven from 5 g corpses. Males feed the larvae as often as females do, and larvae raised by males alone are not significantly different in weight from larvae raised by females alone or by both parents. Males which cared for a succession of broods in the laboratory did not differ significantly in median lifespan from males which were removed from their corpses after eggs had been laid. Non-caring males weighed significantly more than caring males over a sequence of corpses, but the caring males did not differ significantly in weight from non-breeding controls.     

Handling and Survivorship of African Wild Dog (Lycaon pictus) in Five Ecosystems

Populations of the African wild dog, Lycaon pictus have declined throughout their range in sub-Saharan Africa during the last 20 years. It has been hypothesized that handling of wild dogs led to local extinction of a study population in the Serengeti-Masai Mara ecosystem. In this paper we compare rates of mortality and disappearance in handled ( n = 305) versus unhandled ( n = 135) dogs to test the hypothesis that handling leads to increased mortality of Lycaon . We examine data from five ecosystems in which Lycaon have been handled. Our data show that there is no effect of handling on the longevity of Lycaon in any ecosystem studied. Given these data, a more parsimonious explanation of the decimation of the Serengeti-Mara Lycaon population would be that disease alone was responsible for the population collapse and that researcher handling of Lycaon was correlated with, but not causal to, this mortality.     

Male philopatry,extra-pack copulations and inbreeding avoidance in Ethiopian wolves (Canis simensis)

Monogamous pairings have been regarded as the fundamental social unit in all canid species, including those living in packs. In Ethiopian wolves, however, habitat saturation limits dispersal, which raises the question of whether they avoid inbreeding and, if so, by what mechanism. In two study areas Ethiopian wolf packs had stable memberships. Each pack comprised two to eight adult males, one to three adult females, including a clear-cut dominant individual of each sex, together with one to six yearlings and up to six pups (n = 9 packs). Males remained in their natal packs, apparently throughout their lives. Some females also failed to disperse while others dispersed in their second or third year and became floaters. Dominant females monopolized breeding, and were succeeded either by their most dominant daughters (three cases) or by floaters (two cases). In the former case there is potential for incest; however, 70% of 30 copulations observed were between the dominant female of one pack and a male from an adjoining pack. In Ethiopian wolves, under conditions where dispersal is constrained and the potential for inbreeding is high, extra-pack matings (and associated multiple paternity) result in outbreeding. We raise the possibility that extra-pair copulations may be widespread in canid societies and that the monogamy supposedly fundamental to the family may be more sociological than genetic. Received: 15 October 1994/Accepted after revision: 9 December 1995     

Male quality,reproduction, and survival in the great tit (Parus major)

Summary Three possible measures of male quality (social dominance, song, and size), reproduction, and survival were studied in a single population of great tits. Winter dominance position on a feeder was related to strophe length (number of phrases per strophe), inversely related to positive drift (decrease of the singing rate of the phrases in a strophe), but not related to song repertoire size. Neither winter dominance position nor song were related to size (wing length, tarso-metatarsus length, weight).Singing capacity was not correlated with individual reproductive success in a single breeding season, using a rather limited data set. However, better singers (males which sing longer strophes, show less positive drift, and have larger song repertoires) survived better and had a higher individual lifetime reproductive success (on the basis of a male's recruited offspring of all breeding seasons). Our results show that there exist measurable differences whereby birds that are dominant in winter sing better, survive longer, and produce more surviving offspring during their life time. We suggest that differences in male quality are the common cause (direct and indirect) of all these effects.     

Habitat quality and heterogeneity influence distribution and behavior in African buffalo (Syncerus caffer)

Top-down effects of predators on prey behavior and population dynamics have been extensively studied. However, some populations of very large herbivores appear to be regulated primarily from the bottom up. Given the importance of food resources to these large herbivores, it is reasonable to expect that forage heterogeneity (variation in quality and quantity) affects individual and group behaviors as well as distribution on the landscape. Forage heterogeneity is often strongly driven by underlying soils, so substrate characteristics may indirectly drive herbivore behavior and distribution. Forage heterogeneity may further interact with predation risk to influence prey behavior and distribution. Here we examine differences in spatial distribution, home range size, and grouping behaviors of African buffalo as they relate to geologic substrate (granite and basalt) and variation in food quality and quantity. In this study, we use satellite imagery, forage quantity data, and three years of radio-tracking data to assess how forage quality, quantity, and heterogeneity affect the distribution and individual and herd behavior of African buffalo. We found that buffalo in an overall poorer foraging environment keyed-in on exceptionally high-quality areas, whereas those foraging in a more uniform, higher-quality area used areas of below-average quality. Buffalo foraging in the poorer-quality environment had smaller home range sizes, were in smaller groups, and tended to be farther from water sources than those foraging in the higher-quality environment. These differences may be due to buffalo creating or maintaining nutrient hotspots (small, high-quality foraging areas) in otherwise low-quality foraging areas, and the location of these hotspots may in part be determined by patterns of predation risk.     

Risk assessment in Nile tilapia (Oreochromis niloticus) and African mud catfish (Clarias gariepinus) exposed to cassava effluent

Toxicity of cassava effluent in water on aquatic lives was examined via acute and chronic bioassay experiments on Clarias gariepinus (I) and Oreochromis niloticus (II) under laboratory conditions, using standard procedures. The effluent cyanide concentration exceeded the WHO limit for wastewater. Fish body weights and haematological parameters (HMP) significantly decreased with increasing effluent concentration at p≤0.05. Toxicity on HMP for the respective (I) and (II) varied from 5.4 to 52.8; 4.8 to 51.9% for packed cell volume, 4.1 to 43.9; 5.3 to 64.0% for red blood count, 0.0 to 15.7; 0.0 to 61.4% for white blood count, 3.6 to 45.9; 5.2 to 49.5% for haemoglobin, 11.6 to 71.9; 28.4 to 63.8% for total protein, 11.5 to 75.5; 15.0 to 58.2% for albumin and 11.8 to 75.0; 46 to 83.9% for globulin. Acute exposure yielded 96-h LC₅₀ values of 0.45% for (I) and 0.25% for (II) and chronic exposure caused reduced growth and poor blood quality.     

Digit ratio (2D:4D) and dominance rank in female rhesus macaques (Macaca mulatta)

Female rhesus macaques exhibit matrilineal dominance structures, and high dominance rank confers fitness benefits across a lifetime and across generations. Rank effects are “inherited” through social processes that are well understood; however, biological mechanisms that might impact these processes are not well known. Recently, it has been shown that prenatal androgens appear to be implicated in supporting dominance rank hierarchies in some mammals. In humans, interindividual differences in the second (index) to fourth (ring) digit ratio (2D:4D) have been linked indirectly to variation in prenatal androgens, with low 2D:4D in both sexes associated with higher inferred prenatal androgen effects. 2D:4D has also been related to dominant social behavior and has been shown to co-vary with social systems across nonhuman primate species. Here, we investigate how 2D:4D co-varies with socially inherited dominance rank in female rhesus macaques. Low 2D:4D was associated with higher-ranking females, while higher 2D:4D was associated with lower-ranking females. Similar relationships were also shown between ranked families within matrilines. This is the first study to show such a relationship between 2D:4D and dominance rank in a nonhuman primate and suggests that prenatal androgen effects could be involved in the maintenance of dominance rank in female cercopithecine primates.