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.     

Rothschild’s giraffe Giraffa camelopardalis rothschildi (Linnaeus, 1758) in East Africa: A review of population trends,taxonomy and conservation status

Giraffe populations have suffered a 40% decline in the past thirty years, making them a new priority for conservation and there are considerable uncertainty and disagreement over the taxonomic classification of giraffes. Consequently, there has never been a more critical time to fully understand the global population size and distribution of all giraffe subspecies. The Rothschild's giraffe Giraffa camelopardalis rothschildi (Linnaeus, 1758) is arguably one of the most imperilled giraffe subspecies. Once widespread across southern Sudan, Uganda and Kenya, the Rothschild's giraffe is now confined to a few, isolated and enclosed populations throughout Kenya and Uganda, with only one natural population remaining. Information about Rothschild's giraffe population size, distribution, and conservation history is patchy, and confined to recondite or inaccessible resources. Here, I present a review of taxonomy, distribution, conservation status and population size of the Rothschild's giraffe and present the most recent estimate of population sizes based on direct census data. Although the Latin name affixed to Rothschild's giraffe is subject to discussion, continued reference to this subspecies as G. c. rothschildi is strongly recommended.     

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.     

Leadership of herd progressions in the Thornicroft's giraffe of Zambia

In cohesive social groups, travel progressions are often led by dominant or older individuals, but the leadership traits of individuals residing in flexible social systems are poorly known. Giraffe reside in herds characterized by fission–fusion dynamics frequently mediated by kinship. We analyzed 41 years (1971–2012) of longitudinal data collected from a community of Thornicroft's giraffe (Giraffa camelopardalis thornicrofti) living around South Luangwa National Park, Zambia, to assess the characteristics of herd leaders. Movement of giraffe in a single file progression was not associated with either season or time of day, but progressions were significantly more likely to occur when giraffe traveled in open areas. The oldest female in a herd was significantly more likely to be at the front position than expected, occupying the leadership niche on 79% of observations. We reason that matriarchal leadership in giraffe, as in African elephants, Loxodonta africana, is associated with resource learning. Giraffe societies are constructed on a heretofore unrecognized foundation that integrates relatedness and familiarity with matriarchal leadership in herd movement.     

Applying computer‐aided photo‐identification to messy datasets: a case study of Thornicroft's giraffe (Giraffa camelopardalis thornicrofti)

Digital photography enables researchers to rapidly compile large quantities of data from individually identifiable animals, and computer software improves the management of such large datasets while aiding the identification process. Wild‐ID software has performed well with uniform datasets controlling for angle and portion of the animal photographed; however, few datasets are collected under such controlled conditions. We examined the effectiveness of Wild‐ID in identifying individual Thornicroft's giraffe from a dataset of photographs (n = 552) collected opportunistically in the Luangwa Valley, Zambia from March to October 2009. We assessed the programme's accuracy in correctly identifying individuals and the effect of five image quality factors on identification success: blurriness, background type and complexity, amount of sky and the presence of other giraffe. The programme correctly identified individuals in 71.6% of photographs. Background complexity was the only significant variable affecting identification success and removing background imagery reduced identification error by 52.8% (from 28.4 to 13.4%). Our results indicate higher levels of error than previously reported for Wild‐ID. However, they also suggest the programme is an effective tool for quickly identifying individuals in large field datasets, especially if photograph backgrounds are removed beforehand and postanalysis visual verification is performed.     

Updated geographic range maps for giraffe,Giraffa spp., throughout sub‐Saharan Africa,and implications of changing distributions for conservation

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Seasonal and geographical influences on the feeding ecology of giraffes in the Luangwa Valley,Zambia: 1973–2014

Obtaining longitudinal data about the feeding ecology of long‐lived iteroparous mammals is rare, but enhances our understanding of how the environment influences niche breadth and dietary diversity within a species. We analysed forty years of feeding records obtained from a population of Thornicroft's giraffes (Giraffa camelopardalis thornicrofti) living in the Luangwa Valley, Zambia. Giraffes are browsers that have been reported to feed primarily upon Acacia leaves, but their feeding ecology in some locations conflict with this interpretation. Giraffes in the Luangwa Valley fed on 93 identified plant species, but only a few contributed to the bulk of the diet. Niche breadth was quite large (Shannon‐Weiner Diversity Index H′ = 3.699) and about 13% more diverse during the dry, than wet, season. Key species eaten during the dry season were very consistent across decades, with Kigelia africana and Capparis tomentosa prominent at this time. The evolutionary ecology of giraffes has probably benefitted from a foraging strategy that includes a variable and high‐quality diet during the hot, dry season, when feeding pressures are greatest. Giraffe feeding ecology has evolved in conjunction with their physiology, anatomy and morphology, resulting in an animal that is well adapted for survival in an arid environment.     

Population structure and spatial ecology of Kordofan giraffe in Garamba National Park,Democratic Republic of Congo

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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.     

Population structure and phylogeography of the Gentoo Penguin (Pygoscelis papua) across the Scotia Arc

Climate change, fisheries' pressure on penguin prey, and direct human disturbance of wildlife have all been implicated in causing large shifts in the abundance and distribution of penguins in the Southern Ocean. Without mark‐recapture studies, understanding how colonies form and, by extension, how ranges shift is challenging. Genetic studies, particularly focused on newly established colonies, provide a snapshot of colonization and can reveal the extent to which shifts in abundance and occupancy result from changes in demographic rates (e.g., reproduction and survival) or migration among suitable patches of habitat. Here, we describe the population structure of a colonial seabird breeding across a large latitudinal range in the Southern Ocean. Using multilocus microsatellite genotype data from 510 Gentoo penguin (Pygoscelis papua) individuals from 14 colonies along the Scotia Arc and Antarctic Peninsula, together with mitochondrial DNA data, we find strong genetic differentiation between colonies north and south of the Polar Front, that coincides geographically with the taxonomic boundary separating the subspecies P. p. papua and P. p. ellsworthii. Using a discrete Bayesian phylogeographic approach, we show that southern Gentoos expanded from a possible glacial refuge in the center of their current range, colonizing regions to the north and south through rare, long‐distance dispersal. Our findings show that this dispersal is important for new colony foundation and range expansion in a seabird species that ordinarily exhibits high levels of natal philopatry, though persistent oceanographic features serve as barriers to movement.     

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.     

Sharp genetic discontinuity across a unimodal Heliconius hybrid zone

Hybrid zones are powerful natural systems to study evolutionary processes to gain an understanding of adaptation and speciation. In the Cauca Valley (Colombia), two butterfly races, Heliconius cydno cydnides and Heliconius cydno weymeri, meet and hybridize. We characterized this hybrid zone using a combination of mitochondrial DNA (mtDNA) sequences, amplified fragment length polymorphisms (AFLPs), microsatellites and sequences for nuclear loci within and outside of the genomic regions that cause differences in wing colour pattern. The hybrid zone is largely composed of individuals of mixed ancestry. However, there is strong genetic discontinuity between the hybridizing races in mtDNA and, to a lesser extent, in all nuclear markers surveyed. The mtDNA clustering of H. c. cydnides with the H. cydno race from the Magdalena Valley and H. c. weymeri with the H. cydno race from the pacific coast suggests that H. c. cydnides colonized the Cauca Valley from the north, whereas H. c. weymeri did so by crossing the Andes in the southern part, implying a secondary contact origin. Colonization of the valley by H. cydno was accompanied by mimicry shift. Strong ecological isolation, driven by locally adaptive differences in mimetic wing patterns, is playing an important role in maintaining the hybrid zone. However, selection on wing pattern alone is not sufficient to explain the genetic discontinuity observed. There is evidence for differences in male mating preference, but the contribution of additional barriers needs further investigation. Overall, our results support the idea that speciation is a cumulative process, where the combination of multiple isolation barriers, combined with major phenotypic differences, facilitates population divergence in face of gene flow.     

Genetic divergence between the South Korean and Mongolian populations of the dung beetle,Gymnopleurus mopsus (Coleoptera: Scarabaeidae) based on mitochondrial cytochrome c oxidase subunit I (COI) gene sequences

The locally extinct dung beetle, Gymnopleurus mopsus Pallas, 1781 (Coleoptera: Scarabaeidae), has not been found in South Korea since the 1970s. This research was conducted to understand the genetic divergence between the South Korean and Mongolian populations of G. mopsus as a part of its reintroduction program in South Korea. The genetic distance and diversity were determined using the mitochondrial cytochrome c oxidase subunit I (COI) gene sequence (658 bp ) corresponding to the DNA barcode region. The mitochondrial COI gene sequences of 64 individuals of G. mopsus collected in South Korea (7 individuals) and Mongolia (57 individuals) showed a relatively high level of genetic diversity (nucleotide diversity, 0.0078 ± 0.0007; Haplotype diversity, 0.965 ± 0.017). The genetic distances between the South Korean and Mongolian populations lay within the intraspecific level and the phylogenetic reconstruction using the neighbor‐joining (NJ) method showed that all individuals belonged to a single clade. This result indicates that the current Mongolian population of G. mopsus is a good candidate source population to restore the locally extinct population of the species in South Korea.     

Distribution of Lilian's Lovebirds in Malawi

Lilian's Lovebird Agapornis lilianae is regarded as a mopane Colophospermum mopane woodland specialist. Its global population is sparse and is spread along the Zambezi valley with little known about its current distribution and status. We explored the current distribution of Lilian's Lovebird in Malawi, and in particular focused on the distribution and habitat associations of the largest resident population in Liwonde National Park (LNP). Local birders and tourist guides provided distribution information from across Malawi. Transect walks were conducted to collect data in LNP. Five new atlas records are reported; three were within 40–56 km of the LNP population and two were over 150 km south and north of LNP, respectively. One of the sites is about 66 km from the Lilian's Lovebird population in Luangwa Valley, Zambia. New national records were evidence of the importance of seasonal movements to the species. Lilian's Lovebirds occurred throughout LNP with the highest abundance in the central section. Seasonal movements to areas outside the park were also recorded. A variety of vegetation types were used by the lovebirds, but the strongest habitat associations were with seasonally wet grasslands and not mopane woodlands as would be expected. Thus, conservation efforts should also include these other habitats.     

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.     

Microsatellites from the vairone Leuciscus souffia (Pisces: Cyprinidae) and their application to closely related species

Eleven microsatellites were isolated from the vairone Leuciscus souffia (Risso 1826), an endangered fish that inhabits river systems in and around the Alps in Europe. The level of genetic diversity was assessed in 29 individuals of the subspecies L. s. souffia, and their variability was further estimated in seven individuals of a different subspecies, L. s. muticellus. Eight of these microsatellite loci were also applied to seven closely related cyprinid species. Availability of the reported microsatellite loci will facilitate the investigation of population genetic structure of these species with applications for the development of conservation strategies and phylogeographical approaches.     

Population genetic structure,introgression, and hybridization in the genus Rhizophora along the Brazilian coast

Mangrove plants comprise plants with similar ecological features that have enabled them to adapt to life between the sea and the land. Within a geographic region, different mangrove species share not only similar adaptations but also similar genetic structure patterns. Along the eastern coast of South America, there is a subdivision between the populations north and south of the continent's northeastern extremity. Here, we aimed to test for this north‐south genetic structure in Rhizophora mangle, a dominant mangrove plant in the Western Hemisphere. Additionally, we aimed to study the relationships between R. mangle, R. racemosa, and R. × harrisonii and to test for evidence of hybridization and introgression. Our results confirmed the north‐south genetic structure pattern in R. mangle and revealed a less abrupt genetic break in the northern population than those observed in Avicennia species, another dominant and widespread mangrove genus in the Western Hemisphere. These results are consistent with the role of oceanic currents influencing sea‐dispersed plants and differences between Avicennia and Rhizophora propagules in longevity and establishment time. We also observed that introgression and hybridization are relevant biological processes in the northeastern coast of South America and that they are likely asymmetric toward R. mangle, suggesting that adaptation might be a process maintaining this hybrid zone.     

Phylogeography of Chinese house mice (Mus musculus musculus/castaneus): distribution,routes of colonization and geographic regions of hybridization

House mice (Mus musculus) are human commensals and have served as a primary model in biomedical, ecological and evolutionary research. Although there is detailed knowledge of the biogeography of house mice in Europe, little is known of the history of house mice in China, despite the fact that China encompasses an enormous portion of their range. In the present study, 535 house mice caught from 29 localities in China were studied by sequencing the mitochondrial D‐loop and genotyping 10 nuclear microsatellite markers distributed on 10 chromosomes. Phylogenetic analyses revealed two evolutionary lineages corresponding to Mus musculus castaneus and Mus musculus musculus in the south and north, respectively, with the Yangtze River approximately representing the boundary. More detailed analyses combining published sequence data from mice sampled in neighbouring countries revealed the migration routes of the two subspecies into China: M. m. castaneus appeared to have migrated through a southern route (Yunnan and Guangxi), whereas M. m. musculus entered China from Kazakhstan through the north‐west border (Xinjiang). Bayesian analysis of mitochondrial sequences indicated rapid population expansions in both subspecies, approximately 4650–9300 and 7150–14 300 years ago for M. m. castaneus and M. m. musculus, respectively. Interestingly, the migration routes of Chinese house mice coincide with the colonization routes of modern humans into China, and the expansion times of house mice are consistent with the development of agriculture in southern and northern China, respectively. Finally, our study confirmed the existence of a hybrid zone between M. m. castaneus and M. m. musculus in China. Further study of this hybrid zone will provide a useful counterpart to the well‐studied hybrid zone between M. m. musculus and Mus musculus domesticus in central Europe.     

Dual influence of terrestrial and marine historical processes on the phylogeography of the Brazilian intertidal red alga Gracilaria caudata

In this study, we explored how past terrestrial and marine climate changes have interacted to shape the phylogeographic patterns of the intertidal red seaweed Gracilaria caudata, an economically important species exploited for agar production in the Brazilian north‐east. Seven sites were sampled along the north‐east tropical and south‐east sub‐tropical Brazilian coast. The genetic diversity and structure of G. caudata was inferred using a combination of mitochondrial (COI and cox2‐3), chloroplast (rbcL) and 15 nuclear microsatellite markers. A remarkable congruence between nuclear, mitochondrial and chloroplast data revealed clear separation between the north‐east (from 03° S to 08° S) and the south‐east (from 20° S to 23° S) coast of Brazil. These two clades differ in their demographic histories, with signatures of recent demographic expansions in the north‐east and divergent populations in the south‐east, suggesting the maintenance of several refugia during the last glacial maximum due to sea‐level rise and fall. The Bahia region (around 12° S) occupies an intermediate position between both clades. Microsatellites and mtDNA markers showed additional levels of genetic structure within each sampled site located south of Bahia. The separation between the two main groups in G. caudata is likely recent, probably occurring during the Quaternary glacial cycles. The genetic breaks are concordant with (i) those separating terrestrial refugia, (ii) major river outflows and (iii) frontiers between tropical and subtropical regions. Taken together with previously published eco‐physiological studies that showed differences in the physiological performance of the strains from distinct locations, these results suggest that the divergent clades in G. caudata correspond to distinct ecotypes in the process of incipient speciation and thus should be considered for the management policy of this commercially important species.     

7. SHORT NOTES FROM MEMBERS

Cooper, J., Ross, G. J. B. &; Shaughnessy, P. D. 1978. Seasonal and spatial distribution of Rockhopper Penguins ashore in South Africa. Ostrich 49:40-44. There are 30 records of Rockhopper Penguins Eudyptes chrysocome ashore in South Africa. Both the southern subspecies E.c. chrysocome, and the northern subspecies E.c. moseleyi have been recorded. The northern subspecies has occurred more frequently. Most records are of moulting juveniles in January and February. Records of adult birds are more scattered throughout the year. Rockhopper Penguins in South Africa have been recorded only south of 30S. Birds of the northern subspecies probably originate from South Atlantic islands (Tristan da Cunha group and Gough). Birds of the southern subspecies are probably from the Prince Edward Islands.