Carnivory in the common hippopotamus Hippopotamus amphibius: implications for the ecology and epidemiology of anthrax in African landscapes

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Are hippos Africa's most influential megaherbivore? A review of ecosystem engineering by the semi-aquatic common hippopotamus

Megaherbivores perform vital ecosystem engineering roles, and have their last remaining stronghold in Africa. Of Africa's remaining megaherbivores, the common hippopotamus (Hippopotamus amphibius) has received the least scientific and conservation attention, despite how influential their ecosystem engineering activities appear to be. Given the potentially crucial ecosystem engineering influence of hippos, as well as mounting conservation concerns threatening their long-term persistence, a review of the evidence for hippos being ecosystem engineers, and the effects of their engineering, is both timely and necessary. In this review, we assess, (i) aspects of hippo biology that underlie their unique ecosystem engineering potential; (ii) evaluate hippo ecological impacts in terrestrial and aquatic environments; (iii) compare the ecosystem engineering influence of hippos to other extant African megaherbivores; (iv) evaluate factors most critical to hippo conservation and ecosystem engineering; and (v) highlight future research directions and challenges that may yield new insights into the ecological role of hippos, and of megaherbivores more broadly. We find that a variety of key life-history traits determine the hippo's unique influence, including their semi-aquatic lifestyle, large body size, specialised gut anatomy, muzzle structure, small and partially webbed feet, and highly gregarious nature. On land, hippos create grazing lawns that contain distinct plant communities and alter fire spatial extent, which shapes woody plant demographics and might assist in maintaining fire-sensitive riverine vegetation. In water, hippos deposit nutrient-rich dung, stimulating aquatic food chains and altering water chemistry and quality, impacting a host of different organisms. Hippo trampling and wallowing alters geomorphological processes, widening riverbanks, creating new river channels, and forming gullies along well-utilised hippo paths. Taken together, we propose that these myriad impacts combine to make hippos Africa's most influential megaherbivore, specifically because of the high diversity and intensity of their ecological impacts compared with other megaherbivores, and because of their unique capacity to transfer nutrients across ecosystem boundaries, enriching both terrestrial and aquatic ecosystems. Nonetheless, water pollution and extraction for agriculture and industry, erratic rainfall patterns and human–hippo conflict, threaten hippo ecosystem engineering and persistence. Therefore, we encourage greater consideration of the unique role of hippos as ecosystem engineers when considering the functional importance of megafauna in African ecosystems, and increased attention to declining hippo habitat and populations, which if unchecked could change the way in which many African ecosystems function.     

Hippos alter their aggregations to mitigate density-dependent drought effects

Megaherbivores play a critical role in the ecology of African savannas and grasslands. In addition, these systems are forecast to experience more frequent and severe droughts as a product of changes in the global climate. Thus, the continued conservation of megaherbivores and their associated ecosystems will require a better understanding of how megaherbivores respond to drought by shifting their movement, diet and social behaviour. We address this need by investigating the factors affecting changes in the abundance of common hippopotamus (Hippopotamus amphibius; hereafter: ‘hippos’) throughout the six major rivers of Kruger National Park, South Africa, during and following the severe drought of 2015/2016. Specifically, we aimed to understand the role of two environmental characteristics that have relevance to hippos and that changed in response to drought: vegetation condition and the extent of pooled surface water. In addition, we investigated the extent to which pre-drought density affected changes in hippo abundance. Although vegetation and daytime refugia both appeared to influence pre-drought hippo abundance, these factors were less important to the change in hippo abundance related to the drought. Instead, the response to drought was most strongly related to the pre-drought abundance of hippos, where river segments supporting more than 50 individuals prior to the drought in 2015 decreased by more than half on average. Furthermore, we show that the degree of aggregation decreased from 2015 to 2016 because of the drought, but then began to increase again as the rains returned in 2017. Our results suggest that in addition to the large pools that support large aggregations of hippos in typical years, additional smaller pools are likely important for accommodating this drought-induced dispersion. However, maintaining this distribution of pools will likely become more challenging as southern Africa's population and water demands increase.     

Intake, ingesta retention, particle size distribution and digestibility in the hippopotamidae

Although several aspects of the digestive physiology of the hippopotamidae-non-ruminating foregut fermenters-have been described, ingesta kinetics and passage characteristics of these species are not well understood. The most outstanding feature of the hippo digestive physiology reported so far is the very long mean ingesta retention times (MRTs) measured by Foose [Foose, T., 1982. Trophic strategies of ruminant versus nonruminant ungulates. PhD dissertation, University of Chicago, Chicago.]. Since those data had been investigated with animals without water access, we intended to measure MRT in hippos which were allowed to enter water pools during the night. MRT parameters as well as dry matter (DM) digestibility were determined in four common (Hippopotamus amphibius) and four pygmy hippos (Hexaprotodon liberiensis) on two different diets each using cobalt ethylendiamintetraacetate (Co-EDTA) as a fluid, chromium (Cr)-mordanted fibre (<2 mm) as a particle and acid detergent lignin (ADL) as an internal digestibility marker. Four of the animals additionally received cerium (Ce)-mordanted fibres (2-10 mm) as particle markers. Total MRTs for fluids and particles ranged between 20-35 and 48-106 h in the common and between 13-39 and 32-107 h in the pygmy hippos. The difference between fluid and particle retention was greater than usually reported in ruminants. Excretion patterns of the markers differed from those usually observed in ruminants but resembled those reported for macropods (kangaroos), indicating a plug-flow reactor-like physiology in the hippo forestomach (FRST). This finding complements other described similarities between the macropod and the hippo forestomach. The measurements of larger particle retention profiles suggest that in the hippo, larger particles might be excreted either faster or at the same rate as smaller particles, indicating a general difference between ruminants and hippos with respect to differential particle retention. The digestive physiology of hippos is characterised by a generally low food intake, long ingesta retention times and dry matter digestibilities lower than reported in ruminants. Moderate digestibilities in spite of long retention times might be the result of the generally high average ingesta particle size in hippos. The comparatively easy management of pygmy hippos, together with the significant correlations between food intake, MRT and digestibility in the pygmy hippos of this study, recommends this species for further studies on the interplay of these parameters in herbivore digestive physiology.     

Social organization of the spotted hyaena (Crocuta crocuta). I. Demography

A 4-year field study of one group of 60–80 spotted hyaenas (Crocuta crocuta) in Kenya's Masai Mara National Reserve has revealed a form of social system undescribed among carnivores but common among primates. Male juveniles disperse from the natal group around puberty and join a population of nomadic males, wandering widely before settling in a new clan. Adult males frequently leave their adopted clan after a variable period of residence. Females remain in their natal group for life. Thus, the clan comprises several matrilines of related females, while resident adult males originate outside the group. Clans in the Mara tend to be separated by wide buffer zones where prey is scarce and boundary maintenance behaviours are relatively uncommon. Transients are subordinate to residents, but are tolerated by them, in contrast to the more territorial clan system that has been described in Ngorongoro Crater.     

Intraspecific interactions and their effect on habitat utilisation by the praying mantid <Emphasis Type="Italic">Ciulfina biseriata</Emphasis> (Mantodea: Liturgusidae)

Tree trunks are spatially separated habitat patches for the tree-running mantid Ciulfina biseriata (Mantodea: Liturgusidae). This paper investigates how intraspecific interactions may affect spacing patterns in C. biseriata. Many mantid species inhabit tree trunk surfaces, but little is known about how they interact with this type of habitat. Behavioural observations and choice tests investigated the effect of a conspecific on habitat utilisation. Field surveys were used to investigate patterns of movement in the field. The decision to inhabit a tree trunk may be affected by the presence of a conspecific. Whereas adult males seek out females when selecting habitat, females will actively avoid males. Habitat choice decisions are primarily based on aspects of their mating system rather than on competition for space or food resources. The behavioural interactions of C. biseriata are described, including novel observations of same-sex mating behaviour in adult males.     

Distribution and population density of the bushcricket Decticus verrucivorus in a damp-meadow biotope

Summary A population of Decticus verrucivorus (Orthoptera: Tettigoniidae) was studied in the field over two successive years. The majority of insects accumulate only in certain parts of a diverse habitat. In this area the mean population density of adult Decticus verrucivorus was 3.1/100 m². Analysis of nearest neighbour distances shows that males have a clumped distribution in the undisturbed habitat. It is concluded that larval distribution reflects the distribution of oviposition sites but that the distribution of adult males, though also depending on the distribution of natural resources and the climatic conditions, is produced mainly by the acoustic interactions with other males. The distribution of adult females does not appear to be determined by the acoustic behaviour of the males. The role of environmental factors such as the availability of food, humidity and temperature, and of the acoustic communication within this species, is discussed with respect to the problem of the high site fidelity of Decticus verrucivorus.     

Population regulation in the Wood Mouse Apodemus sylvaticus (L.)

Watts 1969 described the population cycle of the Wood Mouse Apodemus sylvaticus as a decline in numbers in spring, a stable period in summer and an increase phase in autumn. Aggression by adult males was considered an important aspect of population regulation limiting male survival in spring and juvenile recruitment in summer. However, recent studies of range behaviour and experimental investigations suggest that male A. sylvaticus are not relevant in population regulation and that the principal regulating factors act on female reproductive success. Female reproductive activity is determined by spatial interactions which, in turn, are determined primarily by the density of females and the proportion of female immigrants in the population. The only impact adult males may have on population size is through infanticide and competition with the females for food.     

Annual cycle of a coypu (myocastor coypus) population: male and female strategies

Male and female strategies were compared for wild–living populations of coypu (Myocastor coypus Molina) censused during two years in marshlands of central–west France. In a habitat composed of a canal network bisecting pasture fields typically 0–5 ha in size, coypu movements were largely restricted to stretches of open water, though males explored further from water than females. Adults occupied home ranges with a mean size of 3–82 ha or 1. 19 km of canal, males on average moving faster over a larger area and using a greater length of canal than females. Movements within partially shared ranges were most positively coordinated between male–female pairs, while male–male interactions were less frequent. Juvenile coypus were largely absent from the late winter population. Synchronization of litters in March–April resulted in a peak of juveniles in early summer. The sex ratio of juveniles and sub–adults reversed from a male bias in summer to a female bias in autumn. Population densities peaked in November at 4 ha, with a significant female bias (1M: 1–6F). In the same month, a neighbouring population at lower density (1 ha^-l after treatment with rodenticide) was significantly male biased (2–7M:lF) and may therefore have been acting as a sink for males dispersing from the high density population. Transients comprised 63% of the population. Females were resident longer than males, 22% of females remaining at least 5 months against 6% of males. Dispersal was recorded only for males aged 6 months or older. The general result, of exploratory behaviour amongst males contrasting with the establishment of longer term home ranges by females, is discussed in terms of male reproductive success and the capacity for females to influence nutrient supply to their litters.     

Behavioural responses of captive hippopotamus(hippopotamus amphibius)in lactation season:Maternal investment and plasticity of infant self-independence

In order to promote hippopotamus management in the captive and ex-situ environment, especially the control of behavioural and physiological status during breeding and lactation seasons, we conducted a preliminary study on behavioural responses of a pair of hippos including both mother and infant in Hangzhou Wildlife Park, China. The study of the captive hippos for about 1-month in the lactation season was carried out during August and September, 2009. The behavioural patterns were identified by all occurrence sampling and instantaneous scanning sampling methods with 5-10 min intervals. As a result, mother-offspring conflicts and interactions did occur throughout the whole study period. Early maternal investment showed a positive trend in activity rhythms (slope = 0.0014, Z = 0.3027, P ＜0.001) and a negative trend (slope = -0.0066, Z = 0.8807, P＜0.001) in territorial occupation of water, all of which supported our hypotheses that the mother hippo might exert less care for the infant and cut down on her own obligations in nursing. For infant self-independence, during the whole lactation season, the primary trends of activities and territorial occupation dynamics of the infant hippo were slightly different from before, judging from linear models (slope = -0.0017, Z = 0. 3309, P＜0.001). However, the frequencies of activities were not stable, especially at around 12 days of age. The trends of territorial occupation (slope = -0.0071, Z= 0. 904, P＜0.001) also showed negative dynamics in water body occupation by the time the infant hippo grew up. The general trend (slope = -0.005, Z = 0.06, P＜0.001) of suckling dynamics was demonstrably negative, with an upwards fluctuation at period 3 (10-15th day). This also illustrated that as the infant developed, the dependency on the mother was reduced at the end of the lactation season. In addition, a sharp decline between P3 and P4 also supported the mother-offspring conflict theory. In general, time budgets of hippos in active behaviour were (31.8±2.1)% for the mother and (32.1 ±2.6)% for the infant. Spatial distributions in water within temporal limitations were (80.1 ±2.7)% for the mother and (81.8±2.7)% for the infant. Behavioural dynamics showed strong synchronous relations between maternal investment and infant independence. Our current short-term investigation proves to be a key in management and conservation of hippopotami during the lactation season.     

Ecological studies on a dragonfly,Nannophya pygmaea ramber (Odonata: Libellulidae) I. seasonal changes of adult population and its distribution in a habitat

Adult population of a dragonfly Nannophya pygmaeaRamber inhabited in a damp ground was investigated with mark-and-recapture method in 1975. The following results about the seasonal changes of the population size and distribution in the habitat were obtained. Adults emerged from late May to mid August. The number of the adults was most abundant in early June, but that of matured males in early July. From the recapture data, the estimate of daily survival rate was 0.82, and the length of immature stage in males was estimated as 5 days or so. Total number of post-teneral adults emerged in the habitat was estimated as about 9,000. The mean crowding-mean density regression method was applied for the analysis of the distribution pattern of the adults. Matured males showed a spaced-out distribution, while females and immatured males distributed themselves rather aggregatively. Such a distribution pattern of the matured male would be attributed to their territorial behaviour. The territorial behaviour was considered to force the matured males to extend the distribution area in July when they were most abundant. From the above-mentioned results and some observations, the meaning of the territoriality in this species was discussed.     

The ecology of the hippopotamus in Liwonde National Park,Malawi: implications for management

The hippopotamus population in Liwonde National Park, Malawi was studied from December 2002 to June 2003. Motorboat censuses along the River Shire counted 640 animals, but because of the large number of hippos in temporary water sources at the time, the true number is probably closer to 950. Marked shifts in hippo distribution from the Shire into temporary water sources occurred as the wet season advanced. Because of the Shire’s year‐round water supply, the hippo population is not regulated by the availability of aquatic refuges, but by food availability. By following feeding tracks, dry season grazing range was estimated to extend 5 km east and 1 km west of the river. Grazing intensity transects and visual estimates indicated only a small portion of this area is suitable for hippo grazing, leading to over‐grazing in suitable areas. Coupled with low primary productivity levels in Liwonde, this means that dry season food competition between hippos and other herbivores is probably high. Recent proposals to raise the Shire’s dry season water level should be considered very carefully, as this will flood late dry season grazing grounds, thereby intensifying grazing competition and increasing grazing pressure in remaining grazing areas, having potentially serious impacts on the animal community.     

Coupling cell growth,proliferation, and death. Hippo weighs in

Four recent papers describe the characterization in Drosophila of Hippo, a serine/threonine kinase of the Sterile 20 (STE20) group, resembling Mst1 and Mst2. Hippo restricts cell growth and cell proliferation, promotes cell death, and interacts with the tumor suppressors Salvador and Warts. This, together with the ability of Mst2 to rescue hippo mutant phenotypes, argues that Mst/Hippo proteins are tumor suppressors.     

Effects of high density on spacing behaviour and reproduction in Akodon azarae: A fencing experiment

We studied the short term spacing behavioural responses of Pampean grassland mouse (Akodon azarae) with regard to population density in four 0.25 ha enclosures (two control and two experimental) in the 2011 breeding season. Based on the hypothesis that A. azarae breeding females exhibit spacing behaviour, and breeding males show a fusion spatial response, we tested the following predictions: (1) home range size and intrasexual overlap degree of females are independent of population density values; (2) at high population density, home range size of males decreases and the intrasexual home range overlap degree increases. To determine if female reproductive success decreases at high population density, we analyzed pregnancy rate, size and weight of litters, and period until fecundation in both low and high enclosure population density. We found that both males and females varied their home range size in relation to population density. Although male home ranges were always bigger than those of females in populations with high density, home range sizes of both sexes decreased. Females kept exclusive home ranges independent of density values meanwhile males decreased home range overlap in high breeding density populations. Although females produced litters of similar size in both treatments, weight of litter, pregnant rate and period until fecundation varied in relation to population density. Our results did not support the hypothesis that at high density females of A. azarae exhibit spacing behaviour neither that males exhibit a fusion spatial response.     

Hostplant-habitat structure and the evolution of butterfly mate-locating behaviour

A hostplant-habitat model is described which explains fundamental differences in mate-locating behaviour between butterfly species, based on a sequence of spatially distinctive activities (habitual patrolling, localized perching, territoriality, lek assembly). It gives equal weight to phylogeny and ecology and is consistent with models for the evolution of territoriality and population structure. Perching repeatedly at characteristic sites, territoriality and lek assembly evolve when resources become associated with distinctive hostplant-habitat structures in which topographic features provide predictable vantage points for acquiring mates. Habitual patrolling is an ancestral behaviour which is suited to widespread and unpredictable resource distributions and to species whose individual apparency detracts from the use of topographic vantage points. Hostplants (emergence sites), nectar and thermoregulation sites typically provide resources and act as precursors to localized perching, territoriality and lek assemblages. But these behaviours fail to evolve in species whose habitats do not include obvious topographic landmarks which provide clear vantage points for males awaiting females and focal points for females searching for males. Where these landmarks occur they result in an increase in receptive female density at those locations compared to the remainder of the habitat.     

A note on group sizes of oribi (Ourebia ourebi, Zimmermann, 1783) from two contrasting sites in Zambia, with and without predation

The oribi (Ourebia ourebi, Zimmermann, 1783) is a small species of antelope widely represented across open grasslands of sub‐Saharan Africa. Although largely territorial, differences in group size and mating systems (monogamy/polygamy), have been linked to habitat conditions and population density. At high population densities, additional males are recruited to assist in territorial defence. Other factors which might impact on group size, include predation threat, especially at low population densities. In this study (1995–98), we recorded group sizes of oribi across two contrasting areas (c. 300 km² each) of Zambia with (Kafue), and without (Bangweulu), large vertebrate predators. A total of 412 groups was recorded across the two sites, with mean sizes of 2.31 (n = 217) at Bangweulu and 2.33 (n = 195) at Kafue. The modal group size was two throughout (range 1–6). Groups were more variable in size and sex composition at Kafue than at Bangweulu, but there was no significant difference in estimates of population density between sites. Mean estimates were 2.02 and 1.90 km^?2 for Bangweulu and Kafue, respectively. Sex ratios (males : females) were biased towards females (1 : 1.72 and 1 : 3.55, respectively) at both sites. Adult males were particularly scarce at Kafue. However, values of density and group size, were both unexceptional for the species. Whilst a modal group size of two is consistent with a monogamous mating system, variations in group composition and size were more difficult to explain. Adult females appeared to retain or recruit additional animals to groups, irrespective of the needs of territorial defence. This suggests that vigilance is an important function of groups on large open plains, especially in the presence of predators. By extension, reversed sexual dimorphism in oribi could reflect increased selection for vigilance duties in females. Predators had no apparent effect on population density, but oribi groups were more variable in size and sex composition in their presence. Adult males may be particularly vulnerable to predation when defending territorial borders at low population densities. However, further work is needed to define group dynamics in this species and to identify causes of mortality amongst adult males.     

Spring dispersal and agonistic behaviour of the Red-legged partridge (Alectoris rufa)

Spring behaviour, dispersal and population changes were studied in two populations of individually marked Red-legged partridges.
Young males took up home ranges close to their natal site but young females dispersed away. The incidence of agonistic behaviour between males, the extent of the spring decline in population density and the distance dispersed by males between the winter and breeding ranges were less in an area with more abundant nesting cover.
Half the neighbours of a typical male came from the same winter social group. These groups were based on family parties. Males from the same winter group showed less frequent agonistic behaviour to each other and had more overlapped home ranges than those from different groups.     

Do interspecific interactions between females drive shifts in habitat use? A test using the lizards Anolis carolinensis and A. sagrei

An important goal in evolutionary ecology is to understand how and why coexisting closely related species partition habitat among themselves. Although studies of interspecific interactions typically focus on males, interactions between females may also play an important role in shaping habitat use within multi‐species communities. The green anole (Anolis carolinensis) exhibits a wide range of habitat use in south‐eastern Louisiana, but its observed habitat use is restricted and altered in areas where it occurs with the introduced Anolis sagrei. We staged interactions between these two species in the laboratory to test the hypothesis that A. sagrei dominate A. carolinensis in contests over shared habitat. We examined whether species identity, bite force, dewlap size, and body size affected the outcome of interspecific interactions between both males and females, and tested the prediction that bite force and size would be the most important determinants of interaction outcomes in both sexes. In male interspecific interactions, we found that individuals with relatively larger dewlaps tended to score higher on aggressive behaviours regardless of species identity, and that interactions consisted of signalling and rarely escalated to physical combat. However, we found that A. sagrei females achieved higher aggressive scores than A. carolinensis females in almost all cases, lending support to the notion that female interspecific behaviour is probably more important than male behaviour in driving changes in habitat use. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 843–851.     

Oxygen cycle in a hippo pool, Serengeti National Park, Tanzania

The vertical distribution of temperature and dissolved oxygen was sampled over 24 h in a pond of the Seronera River inhabited by hippos. The waters were very turbid (visibility < 2 cm) because of the high turbidity from animals trampling sediment and a permanent surface algal bloom sustained by faecal matter. Direct solar heating was restricted to the top few centimetres. This resulted in a strong thermal and density stratification inhibiting aeration of the water column. Waters at mid-depth were aerated only when hippos stirred the water. Anoxic conditions were common in bottom water, these were occasionally ventilated in daytime by mixing due to bottom heating from decaying organic matter and at night by convective cooling. Poor water quality in hippo pools may affect wildlife.