Genetic structure of two populations of the Namibian giraffe,Giraffa camelopardalis angolensis

Two geographically distinct populations of giraffe (Giraffa camelopardalis) were sampled for this study, the northern Namib Desert and Etosha National Park. Population genetic parameters and relationships within subpopulations were estimated to better understand the genetic architecture of this isolated subspecies. Gene flow between the geographically separated populations can be attributed to recent translocation of giraffe between the two populations. Inbreeding estimates in the six subpopulations studied were low though we found evidence that genetic drift may be affecting the genetic diversity of the isolated populations in northern Namibia. Population dynamics of the sampling locations was inferred with relationship coefficient analyses. Recent molecular systematics of the Namibian giraffe populations indicates that they are distinct from the subspecies Giraffa camelopardalis giraffa and classified as G. c. angolensis. Based on genetic analyses, these giraffe populations of northern Namibia, the desert‐dwelling giraffe and those protected in Etosha National Park, are a distinct subspecies from that previously assumed; thus we add data on G. c. angolensis to our scientific knowledge of this giraffe of southern Africa.     

A review of the social behaviour of the giraffe Giraffa camelopardalis: a misunderstood but socially complex species

1. Until around 2000, giraffe Giraffa camelopardalis were believed to have no social structure. Despite a resurgence of interest in giraffe behaviour in around 2010, most studies are of isolated populations, making it difficult to draw general conclusions. Although it is now well established that giraffe social organisation is non-random, there is little consensus as to what influences preferred and avoided associations or the underpinning mechanisms.
2. We test two hypotheses: first, giraffe have a complex cooperative social system, exhibited by 1) stable groups of females, 2) offspring that stay in their natal group for part or all of their lives, 3) support by non-mothers in rearing young, and 4) non-reproductive females in the group; and second, giraffe form matrilineal societies, evidenced by 1) male dispersal, 2) female philopatry, 3) assistance in raising or protecting offspring, and 4) individual benefits gained from social foraging.
3. We reviewed 404 papers on giraffe behaviour and social organisation; captive studies were included where they supplemented information from free-living populations.
4. We show that giraffe exhibit many of the features typical of mammals with complex cooperative social systems and matrilineal societies. However, the social complexity hypothesis posits that such species also require complex communication systems to regulate interactions and relations among group members; giraffe communication systems are poorly understood.
5. Quantifying the fitness and survival benefits of the giraffe’s social organisation is necessary to ensure its long-term survival. Giraffe numbers have declined by 40% since 1985, they have been declared extinct in seven (possibly nine) countries and are listed as Vulnerable by the International Union for Conservation of Nature. We identify research areas that will advance our understanding of giraffe behaviour and conservation requirements.

     

Hierarchical foraging by giraffe in a heterogeneous savannah,Tanzania

Understanding foraging decisions made by wildlife at different spatio‐temporal scales is important for wildlife management and conservation. We tested whether foraging decisions by Masai giraffe (Giraffa camelopardalis tippelskirchi Matschie) differed with scales; habitat, patch and tree in a heterogeneous savannah. We collected data from Arusha National Park, Tanzania, in March–May and August–October 2013. Visual observations were used to collect data on foraging behaviour. Measurements of tree height and stem height and scoring of accumulated browsing were made in 133 patches around trees where the giraffes had been seen browsing, and in a corresponding number of available patches. Giraffes selected Acacia shrub and Dodonea shrub habitats but not for water availability or predator avoidance. For patch use, giraffe selected high quantity of Acacia xanthophloea and Dodonea viscosa. Between plant species, A. xanthophloea was the most preferred and within plant species, tree quality was enhanced by tree height and high score of accumulated browsing. Generally, giraffes selected for A. xanthophloea at all scales.     

Herd composition,kinship and fission–fusion social dynamics among wild giraffe

A variety of social systems have evolved as a consequence of competition and cooperation among individuals. Giraffe (Giraffa camelopardalis sp.) societies are an anomaly because the dearth of long‐term data has produced two polar perspectives: a loose amalgamation of non‐bonded individuals that sometimes coalesce into a herd and a structured social system with a fission–fusion process modifying herd composition within a community. We analysed 34 years of data collected from a population of Thornicroft's giraffe (G. c. thornicrofti, Lydekker 1911) residing in South Luangwa, Zambia, to establish the nature of giraffe society. Our sample consisted of 52 individually recognized animals. We found that giraffe herd composition is based upon long‐term social associations that often reflect kinship, with close relatives significantly more likely than non‐relatives to establish herds. Mother/offspring dyads had the strongest associations, which persisted for years. Giraffe live in a complex society characterized by marked flexibility in herd size, with about 25% of the variance in herd composition owing to kinship and sex. We suggest that giraffe herds share many characteristics of fission–fusion social systems and propose that sophisticated communication systems are a crucial component regulating subgroup dynamics.     

The social organization of giraffes in Niger

The social organization of the last population of giraffes ( Giraffa camelopardalis ) in West Africa was studied between October 1996 and December 1997. Population size increased from 51 to 63 individuals during the study period. Groups were larger during the rainy season (mean group size 9.4) than during the dry season (mean group size 6; U = 4131; P < 0.01). Giraffes did not show strong preferential associations. Group types observed were similar to those expected on the assumption that associations are independent of sex and age. Inter-individual associations were low (mean simple ratio value = 0.1), with an individual associating with only half of its partners of the previous day. Young males interacted more frequently than other classes did. Agonistic interactions occurred mostly among males. Contact interactions (i.e. non-agonistic or sexual) occurred between both sexes and most age classes. Links between local communities, domestic animals, fauna and the environment suggest that environment and development cannot be dissociated in the sustainable exploitation of natural resources.     

Social and demographic influences on the feeding ecology of giraffe in the Luangwa Valley,Zambia: 1973–2014

Body size influences metabolic rate, which impacts feeding ecology. Body mass differs by sex in size‐dimorphic species, such as giraffes, and also by age. Giraffes reside in a fission–fusion social system, which influences feeding ecology due to frequent changes in group size and composition. We analysed 40 years of feeding records collected from a population of Thornicroft's giraffes (Giraffa camelopardalis thornicrofti) living in the Luangwa Valley, Zambia. We examined the influence of herd composition, age and sex on diet. Solitary males and herd males did not differ in diet. Dietary diversity was comparable for females and males, with sex differences in plant species eaten present during the dry season. Age differences in feeding ecology were pronounced, with juveniles often feeding on bushes and smaller trees, while adults tended to feed upon taller trees. Both sexes have evolved foraging strategies that maximize nutrient and energy intake commensurate with their reproductive strategies, with male metabolic requirements sometimes greater, and sometimes less, than that of females. We propose that females are not exchanging foraging efficiency with reproductive tactics by feeding on smaller trees in the open, but are increasing prospects for their calves to survive when confronted with interspecific competition by browsers.     

Female tannin avoidance: a possible explanation for habitat and dietary segregation of giraffes (Giraffa camelopardalis peralta) in Niger

Niger is host to the last free‐roaming herd of G.c. peralta (Giraffa camelopardalis peralta). We examined the foraging preferences of these giraffe in their dry‐season habitats, with the goal of preserving the herd in the regions that they currently inhabit. The current dry‐season habitat comprises two distinct vegetation zones. In both of these zones the giraffe must exist alongside the people of this region. The giraffes exhibit a sexual segregation in their dry‐season habitat selection and forage choices. The females show a strong preference for the intermediate zone (IM) when lactating. The males and pregnant females show a preference for the Dallol Bosso (DB). Nursing cows exhibit an avoidance of tannins. Bulls and non‐nursing cows prefer high protein and high fat forage, while subadults show a strong preference for high protein and carbohydrate contents and moderate tannin levels. Combretum glutinosum is the preferred species for adults of both sexes in the IM. Males and females have strong preferences for both Acacia nilotica and Acacia seyal in the DB. Sub‐adults of both sexes strongly prefer Prosopis africana in the IM. Unlike females, males retain their preference for A. nilotica when in the IM.     

华芬酶-1600对长颈鹿日粮营养物质消化率的影响

在长颈鹿日粮中添加精料量的 0 0‰、 0 8‰、 1 0‰、 1 2‰剂量华芬酶 - 16 0 0 ,采用分期试验法对 3只成年健康长颈鹿进行 4期消化试验。结果表明 ,添加外源酶后 ,长颈鹿对日粮干物质、粗蛋白、粗脂肪、粗纤维的消化率均明显提高 (P <0 0 5 ) ,尤其是粗纤维的消化率 ,以第Ⅳ期 (添加 1 2‰ )最为显著 (P <0 0 5 )。因此 ,推荐添加量为 1 2‰。     

Characterization of 16 microsatellite marker loci in the Maasai giraffe (Giraffa camelopardalis tippelskirchi)

Sixteen polymorphic microsatellite markers with an average allele size of k = 4.3 are identified from a genomic plasmid library constructed for giraffe (Giraffa camelopardalis ssp.). Primer sequences and marker data are reported in tabular form. The markers were screened in a population of 25 Maasai giraffe (G. c. tippelskirchi) collected near the Athi River, Kenya. The average observed heterozygosity for each marker was 0.36 with an average expected heterozygosity of 0.535. Hardy–Weinberg deviations are reported from this population, which is suspected to be inbred. The markers will be used to screen the captive giraffe population for subspecific or hybrid classification.     

Socially Defined Subpopulations Reveal Demographic Variation in a Giraffe Metapopulation

Populations are typically defined as spatially contiguous sets of individuals, but large populations of social species can be composed of discrete social communities that often overlap in space. Masai giraffes (Giraffa camelopardalis tippelskirchi) of Tanzania live in distinct social subpopulations that overlap spatially, enabling us to simultaneously explore environmental and social factors correlated with demographic variation in a metapopulation of >1,400 adult females and calves. We considered statistically distinct communities in the social network as subpopulations and tested for variation among the 10 subpopulations in adult female survival, calf survival, and reproductive rate (calf-to-adult female ratio). We then related variation in demographic rates among subpopulations to differences in vegetation, soil type, proximity to 2 types of human settlements, local giraffe population density, and social metrics of relationship strength and exclusivity among adult females. We did not find any among-subpopulation effects on adult female survival, suggesting adult female survival is buffered against environmental heterogeneity among subpopulations. Variation in calf demographic rates among subpopulations were correlated with vegetation, soils, anthropogenic factors, and giraffe population density but not with adult female relationship metrics, despite substantial spatial overlap. Subpopulations with more dense bushlands in their ranges had lower calf survival probabilities, and those closer to human settlements had higher reproductive rates, possibly because of spatial gradients in natural predation. Reproductive rates were higher in subpopulations with more volcanic soils, and calf survival probabilities were greater in subpopulations with higher local adult female densities, possibly related to higher-quality habitat associated with fertile soils or lower predation risk, or to greater competitive ability. The variation in fitness among subpopulations suggests that giraffes do not move unhindered among resource patches to equalize reproductive success, as expected according to an ideal free distribution. The differences in calf survival and reproductive rates could rather indicate intercommunity differences in competitive ability, perception, learning, or experience. Our approach of comparing demography among spatially overlapping yet distinct socially defined subpopulations provides a biologically meaningful way to quantify environmental and social factors influencing fine-scale demographic variation for social species. © 2021 The Wildlife Society.     

High interannual variation in the diet of a tropical forest frugivore (Hylobates lar)

Frugivores must deal with seasonal changes in fruit availability and changes from year to year, as most species of tropical forest fruiting trees have considerable interannual variation in phenology and many are mast fruiters. We quantified seasonal and interannual changes in the fruit diet in a frugivore and important seed disperser, the white‐handed gibbon, Hylobates lar, in Thailand. We used 40‐d following data during April and May replicated in six consecutive years to study interannual variability in the diet and compared it with seasonal changes measured in monthly samples of the same size collected in three successive years. The 40‐d periods of following also allowed us to measure the decline in dietary similarity with time over a finer scale. We measured fruit diet similarity between replicated 5‐d periods using the percentage overlap (Renkonen's) index and Jaccard's similarity index. Seasonally, average dietary overlap between adjacent months was low, and similarity approached zero after four months. Average rate of decline in similarity exceeded 20 percent per 5‐d period. Variation in fruit species in the diet between years was high and was correlated with interannual variation in fruiting phenology. The strongest correlation occurred in the case of Nephelium melliferum, a highly preferred species that dominated the diet in good fruiting years. It is difficult to separate changes in food species preference from changes in availability from year to year. We devised a relative measure of preference that depends on the degree to which the gibbons rely on prior knowledge to find sources.     

Sociability increases survival of adult female giraffes

Studies increasingly show that social connectedness plays a key role in determining survival, in addition to natural and anthropogenic environmental factors. Few studies, however, integrated social, non-social and demographic data to elucidate what components of an animal''s socio-ecological environment are most important to their survival. Female giraffes (Giraffa camelopardalis) form structured societies with highly dynamic group membership but stable long-term associations. We examined the relative contributions of sociability (relationship strength, gregariousness and betweenness), together with those of the natural (food sources and vegetation types) and anthropogenic environment (distance from human settlements), to adult female giraffe survival. We tested predictions about the influence of sociability and natural and human factors at two social levels: the individual and the social community. Survival was primarily driven by individual- rather than community-level social factors. Gregariousness (the number of other females each individual was observed with on average) was most important in explaining variation in female adult survival, more than other social traits and any natural or anthropogenic environmental factors. For adult female giraffes, grouping with more other females, even as group membership frequently changes, is correlated with better survival, and this sociability appears to be more important than several attributes of their non-social environment.     

The effects of a low-carbohydrate ketogenic diet and a low-fat diet on mood, hunger, and other self-reported symptoms

Objective: To investigate the effects of weight loss diets on mood, food cravings, and other self‐reported symptoms. Research Methods and Procedures: Mood and other symptoms were evaluated by participant self‐report using the Atkins Health Indicator Test (AHIT) in individuals undergoing weight loss following either a low‐carbohydrate, ketogenic diet (LCKD) or a low‐fat diet (LFD). Participants were 119 overweight community volunteers randomized to an LCKD or an LFD. An additional 51 participants who had completed an earlier trial contributed data for the psychometric analyses but were not included in the prospective analyses. Self‐reported symptom levels on seven scales factor‐analytically derived from the AHIT (negative affect, fatigue, somatic symptoms, physical effects of hunger, insomnia, hunger, and stomach problems) were acquired during 12 visits. Results: After adjusting for the change in BMI over the course of the trial, participants experienced significant improvements in most symptoms regardless of diet. Diet group × visit interactions were observed for negative affect [F(9,803) = 2.30, p = 0.015] and hunger [F(9,803) = 3.62, p < 0.0002]. Examination of means indicated that the LCKD group reported less negative affect and hunger, compared with the LFD group. Discussion: Regardless of diet, participants experienced significant improvement in a broad range of symptoms. Symptoms of negative affect and hunger improved to a greater degree in patients following an LCKD compared with those following an LFD. Whether these symptom changes explain the greater short‐term weight loss generally experienced by LCKD followers deserves further research.     

The composition and function of all‐male herds of Thornicroft's giraffe,Giraffa camelopardalis thornicrofti,in Zambia

Temporary all‐male social groups are formed in a number of animal species. We examined 34 years of data collected from 36 male Thornicroft's giraffe in the Luangwa Valley, Zambia, to test a set of predictions related to five possible functions of all‐male herds (predator protection, practicing aggressive skills, prolonging life, nutritional demands and resource learning). We found that all‐male herds were significantly smaller than mixed‐sex herds, usually contained a mature bull, and were not dependent upon season or habitat. Dyadic associations between males in single sex herds were quite weak, with <25% of potential male dyads sighted together in an all‐male herd. Our data are best explained as a resource learning strategy adopted by males to obtain more extensive knowledge about the habitat, including both food and female distribution. However, other benefits in the form of predator protection, dietary intake and sharpening competitive skills for future contests over estrous females also seem to mediate formation of giraffe all‐male groups. We conclude that the primary advantage of roaming in all‐male herds changes during the life history of males.     

Does diet in Celtic Sea fishes reflect prey availability?

Feeding preferences of Celtic Sea fishes were investigated using a database of stomach content records, collected between 1977 and 1994. The diet of cod Gadus morhua, hake Merluccius merluccius, megrim Lepidorhombus whiffiagonis, whiting Merlangius merlangus and saithe Pollachius virens changed markedly as the animals grew larger, and although large predators generally chose larger bodied prey, the variability of prey sizes consumed also increased. Large predators continued to select small, low value, benthic prey (e.g. Callionymus spp. and Trisopterus spp.) which were easier to catch, rather than larger, more energy lucrative pelagic prey (e.g. mackerel Scomber scombrus), even though these pelagic prey‐fishes were nearly always available and were often very abundant. Stock estimates of the International Council for the Exploration of the Sea and U.K. groundfish survey catches were used as indices of prey abundance. Blue‐whiting Micromesistius poutassou and other small pelagic fishes (Argentina spp. and clupeoids) were identified as being particularly important, and were consumed by some predators more often than would be expected given the abundance of these prey in the environment. There was no evidence for density‐dependent feeding by predators on mackerel and only hake exhibited density‐dependent feeding on horse‐mackerel. Hake, cod and megrim consumed more blue‐whiting when this prey was at higher abundance in the environment. In choosing what prey to consume, predators must balance costs and benefits, considering the quality of prey and the energy expended during search, capture and handling.     

Seasonal diet preferences of black rhinoceros in three arid South African National Parks

Black rhinoceros diet and browse availability was investigated in Augrabies Falls (AFNP), Karoo (KRNP) and Vaalbos National Parks (VNP) in South Africa. Rhino tracks were followed and 18,804 standard bite volumes recorded. Browse availability of each plant species was recorded by measuring 14,800 plants and calculating reachable browse volume. The diet comprised 51, 53 and 41 plant species in AFNP, KRNP and VNP, respectively, but three species accounted for more than 65% of the diet in each park, making diet less diverse than available browse in AFNP and KRNP. Browse availability explained 14%, 15% and 52% of diet selection in AFNP, KRNP and VNP, with consumption of most plant species significantly different from availability. A few plant species were so highly preferred that browsing intensities were unsustainable, while some common species were totally rejected. Pressure on the eaten browse averaged 4.4%, 14.5% and 1.3% annually of the volume in AFNP, KRNP and VNP. In the dry season, there was a two to sevenfold increase in browsing pressure on species with actively photosynthesizing tissues. A few key species, including Monechma and Zygophyllum species, were identified as potential early warning indicators of black rhino browsing impact.