Female nile hippopotamus (Hippopotamus amphibius) space use in a naturalistic exhibit

Zoological institutions provide naturalistic exhibits for their animals in order to offer a more appealing look for visitors and give the animal the opportunity to engage in more natural behaviors. Examining space use of the animals in the naturalistic exhibit may aid in the management of these animals and inform future naturalistic exhibit design. The hippopotamus is an amphibious ungulate that spends much of its days in the wild in the water but may be found along the banks of the rivers basking in the sun. Our objective was to determine how captive female hippos utilize their exhibit by examining whether hippos selected for certain areas of a naturalistic exhibit. Scan sample data were collected on a group of nine captive female hippos housed at Disney's Animal Kingdom®. Using ArcView, the data were analyzed to determine distribution of hippos in the exhibit and their utilization of depth categories while in the water. Hippos were found to aggregate in preferred areas of the exhibit, mostly water, and selected most for water depths of 0.6–1.0 m. These results will aid in the understanding of hippopotamus space use and may aid zoological institutions in the design of naturalistic exhibits for hippos. Zoo Biol 31: 129–136, 2012. © 2010 Wiley Periodicals, Inc.     

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

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Social behaviors within a group of captive female Hippopotamus amphibius

Grouping is known to occur in many species of mammals, and the structure of groups can range along a continuum from basic aggregations to complex social systems. Any social patterns that may occur within the group must be determined in order to understand the adaptive nature of the group. Female Hippopotamus amphibius are known to aggregate in the wild, but their social behaviors are still not understood. Our objective was to determine if captive female hippos display social structure within an aggregation by examining their interactions, and if kinship, familiarity, and dominance influence these interactions. Behavioral data, using continuous focal animal sampling and scan sampling, were collected on a group of captive female hippos housed at Disney’s Animal Kingdom and were used to analyze their interactions, association patterns based on kinship and familiarity, and a dominance hierarchy. Our results support the hypothesis that hippos exhibit social patterns due to the attraction to particular individuals. There were more associations between kin than non-kin and also between individuals that were more familiar. Dominance patterns were also found among these hippos. These results may aid in the general understanding of hippopotamus behavior and provide a framework for future research.     

Hind limb myology of the common hippopotamus,Hippopotamus amphibius (Artiodactyla: Hippopotamidae)

Based on morphological traits, hippos have traditionally been classified with pigs and peccaries in the suborder Suiformes. However, molecular data indicate that hippos and cetaceans are sister taxa. This study analyses muscle characters of the common hippo hind limb in order to clarify the phylogenetic relationships and functional anatomy of hippos. Several muscles responsible for propelling the body through water are robust and display extensive fusions, including mm. semimembranosus, semitendinosus, biceps femoris and gluteus superficialis. In addition, common hippos retain long flexor and extensor tendons for each digit, reflecting the fact that all four toes are weight‐bearing. These flexor tendons, together with the well‐developed intrinsic muscles of the pes, serve to adduct the digits, preventing splaying of the toes when walking on soft terrain. Lastly, common hippos retain a number of primitive features, including the presence of m. articularis coxae, a well‐developed m. obturator internus, superficialis and profundus tendons to all digits, mm. flexor digitorum brevis, abductor digiti V, lumbricalis IV, adductores digitorum II and V, and two mm. interossei per digit. Pygmy hippos share these features. Thus, hippopotamids retain muscles that have been lost in the majority of artiodactyls, including other suiforms. These and previously reported findings for the forelimb support the molecular data in indicating an early divergence of the Hippopotamidae from the rest of the Artiodactyla. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 661–682.     

Plant water uptake by hard red winter wheat (Triticum aestivum L.) genotypes at 2°C and low light intensity

Background  

Hard red winter wheat (HRWW; Triticum aestivm L.) plants from genotypes selected in the Northern Great Plains of the U.S. have less tissue water after exposure to cool autumn temperatures than plants from the Southern Great Plains. It is generally assumed that the reduced tissue water content of northern compared to southern cultivars is due to an impedance to water uptake by northern plants as a result of the low autumn temperatures. We hypothesize that if low temperature impedes water uptake then less soil water would be removed by northern than by southern-selected cultivars. This hypothesis was tested by comparing plant water uptake of a northern (FR) and a southern (FS) cultivar in relation to their foliage water content at 2°C.     

Activity in Heodes virgaureae (Lep., Lycaenidae) in relation to air temperature,solar radiation,and time of day

Summary The degree of activity of H. virgaureae in the field is largely dependent on air temperature, solar radiation, and wind velocity. Solar radiation increases body temperature above ambient. The butterfly orientates its back towards the sun and exposes the dorsal surface of the wings. At high temperatures they close the wings thereby minimizing the surface exposed to the sun. The optimal body temperature lies around 35°C as was indicated by laboratory experiments. In cloudy and cool to fairly warm conditions the butterfly is inactive. In sunshine the butterfly basks at low radiation intensities or low air temperatures while feeding (in males also flying) predominates at full sunshine or very high air temperatures (around 30°C). Males fly 5–10 times as much as females. A change from unfavourable to favourable weather is followed by an immediate increase in activity of the butterfly, which enables the butterfly to utilize short periods of sunshine.     

First documentation of flehmen in a common hippopotamus (Hippopotamus amphibius)

A male common hippopotamus (Hippopotamus amphibius) was observed giving a flehmen grimace on 14 occasions in July and August of 1998 at the Denver Zoo. Twelve instances of flehmen occurred after anogenital investigation of the female as she was exiting the pool. The two other instances of flehmen occurred in the terrestrial portion of the enclosure after the male examined samples of the female's freshly voided urine. The frequency of flehmen increased as the female approached estrus. Flehmen in common hippos may be an example of an evolutionary remnant. Zoo Biol 18:415–420, 1999. © 1999 Wiley‐Liss, Inc.     

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.     

Behavioural responses of <Emphasis Type="Italic">ex-situ</Emphasis> captive hippopotamus (<Emphasis Type="Italic">Hippopotamus amphibius</Emphasis>) 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.     

Consequences of dam‐altered thermal regimes for a riverine herbivore's digestive efficiency,growth and vulnerability to predation

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Suppression of ovulation in Nile Hippopotamus (Hippopotamus amphibious) using melengestrol acetate‐treated feed or high dose Depo‐Provera injection

Analysis of fecal progestogen profiles during Depo‐Provera injection (1,200 mg; DEPO, Pfizer Inc., New York, NY), melengestrol acetate (MGA) in feed (2 or 3 mg/head/day), and a combination treatment (DEPO+MGA) are presented for nine captive female Nile hippos housed at Disney's Animal Kingdom in Florida. All tested treatments reduced fecal progestogen elevations successfully to durations consistent with prevention of ovulation for a portion of the treatment period. Percentage of treatment months with suppression of luteal phases indicative of ovulation was maximal for high‐dose MGA (91.7±13.9%) and DEPO+MGA (91.7±20.4%), followed by DEPO injection alone (69.2±13.9%) and low‐dose MGA (57.6±33.2%). Both 1,200 mg DEPO and low‐dose MGA (2.0 mg/day) treatments were insufficient to prevent an apparent seasonal breakthrough of ovarian activity from June–August 2002. Although luteal phases were observed, no females conceived during those months. Overall, in 133.5 treatment months with females housed with an adult male, one female conceived during the transition period between treatments. After cessation of contraceptive treatment, average latency to first normal ovarian cycle was 80.6±19.5 days (range = 22–179 days). Up to 12 months post‐treatment, however, successive cycles were often irregular with evidence of short periods of anovulation and shortened luteal phases in all females monitored. In conclusion, high dose and combination treatments were most successful in preventing progestogen increases indicative of ovulation in hippos. Zoo Biol 26:259–274, 2007. © 2007 Wiley‐Liss, Inc.     

PHOTOSYNTHETIC SENSITIVITY TO TEMPERATURE IN POPULATIONS OF TWO C4 BOUTELOUA (POACEAE) SPECIES NATIVE TO DIFFERENT ALTITUDES

The relationships between photosynthesis, flowering, and growth temperatures were examined experimentally in four populations of the C₄ grass genus Bouteloua. Field-collected plants were grown under two temperature regimes, cool (20 C day/6 C night) and warm (30/16), representative of the extreme populations. Populations collected from the warm climates had significantly lower photosynthetic capacity when grown in the cool chamber relative to the warm chamber, while photosynthetic capacity in the cool climate populations did not differ between the growth conditions. Additionally, exposure to a 2-day cold temperature treatment (10/-2), representative of late-season frosts in high altitude sites, resulted in further reductions in photosynthesis in the warm climate plants, but not in the cool climate plants. This effect was greater for plants grown in the cool growth chamber. Flowering was reduced by 70% in the warm climate plants grown in the cool chamber, and was correlated with photosynthetic inhibition following the short-term cold temperature treatment. These results indicate that genetic differentiation for photosynthetic temperature sensitivity has occurred in the cool climate populations, and that long-term exposure to cool temperatures coupled with short-term relatively extreme low temperatures results in greater photosynthetic inhibition in nontolerant populations.     

Dynamic photo-inhibition and carbon gain in a C4 and a C3 grass native to high latitudes

C₄ plants are rare in the cool climates characteristic of high latitudes and altitudes, perhaps because of an enhanced susceptibility to photo‐inhibition at low temperatures relative to C₃ species. In the present study we tested the hypothesis that low‐temperature photo‐inhibition is more detrimental to carbon gain in the C₄ grass Muhlenbergia glomerata than the C₃ species Calamogrostis Canadensis. These grasses occur together in boreal fens in northern Canada. Plants were grown under cool (14/10 °C day/night) and warm (26/22 °C) temperatures before measurement of the light responses of photosynthesis and chlorophyll fluorescence at different temperatures. Cool growth temperatures led to reduced rates of photosynthesis in M. glomerata at all measurement temperatures, but had a smaller effect on the C₃ species. In both species the amount of xanthophyll cycle pigments increased when plants were grown at 14/10 °C, and in M. glomerata the xanthophyll epoxidation state was greatly reduced. The detrimental effect of low growth temperature on photosynthesis in M. glomerata was almost completely reversed by a 24‐h exposure to the warm‐temperature regime. These data indicate that reversible dynamic photo‐inhibition is a strategy by which C₄ species may tolerate cool climates and overcome the Rubisco limitation that is prevalent at low temperatures in C₄ plants.     

C<Subscript>4</Subscript> grasses in boreal fens: their occurrence in relation to microsite characteristics

C₄ plants are rare in cool climates, an ecological pattern attributable to their poor photosynthetic performance at low temperatures relative to C₃ species. However, some C₄ species are able to persist at high latitudes and high elevations, possibly due to the characteristics of the particular microsites they inhabit in these otherwise unfavourable environments. One such species is Muhlenbergia glomerata, which occurs above 60°N in Canada and is found in the atypical C₄ habitat of boreal fens. In this study, we evaluate how microsite features affect the success of M. glomerata in boreal fens. We surveyed 19 populations across northern Ontario during the summers of 1999 and 2000. The ground coverage by woody vegetation was the most important parameter affecting the presence or absence of M. glomerata. Woody plants covered over 50% of the ground area in plots where M. glomerata is absent, but less than 20% where it is present. The minimum light intensity threshold for the presence of the C₄ species was about 32% of full-sunlight at plant height. Surprisingly, in boreal fens M. glomerata was largely restricted to the wetter moss hollows, rather than occurring on the dry hummocks where its greater water use efficiency might have been advantageous. Woody species dominated the hummocks, but were uncommon in the hollows. In these cool northern climates M. glomerata apparently persists because sufficient periods of temperatures favourable to C₄ photosynthesis occur, but this persistence likely requires some factor that suppresses the woody vegetation.     

Streptomycin mimics the cool temperature response in rice plants

Exogenous application of streptomycin to etiolated seedlings of rice (Oryza sativa L.) during growth in darkness at moderate temperatures induced the same type of chlorosis as that elicited by cool temperatures. A comparison of sensitive (Indica) and tolerant (Japonica) cultivars indicated a close relationship between sensitivity to streptomycin and cool temperatures. Immunoblot (Rubisco LSU, SSU; CF1 complex of H⁺-ATPase; NADPH-protochlorophyllide oxidoreductase) and Northern blot analyses of plastid-encoded genes (16S rRNA; rbcL; rpoB; petB) in the streptomycin-treated sensitive cultivars revealed that the normal etioplast development was specifically inhibited by the antibiotic. Furthermore, the antibiotic did not affect the expression of mitochondrion-encoded genes (18S rRNA; atpA), which are also unaffected by cool temperatures. These result suggest that the effect of the antibiotic is quite similar to that of cool temperatures.     

Prevailing sea surface temperatures inhibit summer reproduction of the kelp Laminaria digitata at Helgoland (North Sea)

The impact of abiotic factors on kelp sporophyte reproduction has rarely been investigated. Laminaria digitata (Hudson) J.V. Lamouroux is one of the few summer fertile Laminaria species worldwide and reproduction is subjected to relatively high water temperatures. We investigated the impact of prevailing summer temperatures (~18°C in August) on the induction of sporangia, meiospore release, and germination at the island of Helgoland (North Sea). At Helgoland, fertile sporophytes are found between April and December with a maximum in late summer. While released meiospore numbers were constant between June and October, germination rates decreased significantly in summer. Short‐term exposure of mature sori to 17°C–22°C induced a significantly higher meiospore release indicating enhancement of sporulation by elevated temperatures. Induction of sporangia on vegetative blade disks was not possible at 20°C, and fertility was only 20% at 18°C–19°C, but it was 100% in cool temperatures of 1°C–10°C. It was shown for the first time in a kelp species that “sporogenesis” is the life‐cycle process with the narrowest temperature window compared to growth or survival of the sporophyte or reproduction, growth, and survival of the gametophyte. We incorporated several parameters (induction time, fertile area, and relative fertility) into a “Reproductive efficiency index.” This indicates that sporogenesis of L. digitata is a cold‐adapted process with an optimum at (5)–10°C. The results show that the population at Helgoland is at its reproduction limit despite the existence of other geographically more southerly located populations.     

Early seedling and seasonal N2-fixing symbiotic activity of two soybean [Glycine max (L.) Merr.] cultivars inoculated with Bradyrhizobium strains of diverse origin

In areas with a short growing season the poor adaptability of soybean [Glycine max Meer. (L.)] to cool soil conditions is considered the primary yield limiting factor. Soybean requires temperatures in the 25 to 30°C range for optimum N₂-fixation and yield. Field studies were conducted in 1990 and 1991 at Montreal, Quebec to determine whether adaptability to cool soil conditions, with respect to earlier symbiosis establishment and function, existed among either Bradyrhizobium strains or soybean genotypes. An early maturing isoline of the soybean cultivar Evans and the cultivar Maple Arrow were inoculated with one of four strains isolated from the cold soils of Hakkaido, northern Japan, or the commercially used strains 532C or USDA110, at two planting dates. Plot biomass and nodulation were assessed at seedling (V2), and flowering(R2) growth stages and harvest maturity. Soybean genotypes did not differ for pre-flowering nodulation or N₂-fixation in the cool spring conditions of the first year. Seasonal N₂-fixation rates were also determined at the final harvest by the N-balance and ¹⁵N-isotope dilution methods. Significantly higher symbiotic activity was found for two of the four Hakkaido strains and was reflected in higher final soybean seed yield and total N₂-fixation for the growing season, as compared to the two commercial strains. Planting 14 days earlier resulted in greater early vegetative and total seasonal N₂ fixation and yield in the second year when soil temperatures were warmer, emphasizing the need for the development of soybean-Bradyrhizobium combinations superior in nodule development and function under cool soil conditions.     

More efficient mastication allows increasing intake without compromising digestibility or necessitating a larger gut: Comparative feeding trials in banteng (Bos javanicus) and pygmy hippopotamus (Hexaprotodon liberiensis)

The digestion of plant material in mammalian herbivores basically depends on the chemical and structural composition of the diet, the mean particle size to which the forage is processed, and the ingesta retention time. These different factors can be influenced by the animal, and they can presumably compensate for each other. The pygmy hippopotamus, a non-ruminating foregut fermenter, has longer mean retention times than ruminants; however hippos do not achieve higher (fibre) digestibilities on comparable diets, which could be due to ineffective mastication. We performed feeding trials with six pygmy hippos (Hexaprotodon liberiensis) and six banteng cattle (Bos javanicus) on a grass diet. As predicted, both species achieved similar dry matter, organic matter, crude protein and gross energy digestibilities. However, neutral and acid detergent fibre digestibility was lower in pygmy hippos. Apparently, in these species, fibre digestibility was more influenced by particle size, which was larger in pygmy hippos compared to banteng, than by retention time. In spite of their higher relative food intake, the banteng in this study did not have greater relative gut fills than the hippos. Ruminants traditionally appear intake-limited when compared to equids, because feed particles above a certain size cannot leave the rumen. But when compared to nonruminating foregut fermenters, rumination seems to free foregut fermenters from an intrinsic food intake limitation. The higher energy intakes and metabolic rates in wild cattle compared to hippos could have life-history consequences, such as a higher relative reproductive rate.     

Takeoff temperatures in Melitaea cinxia butterflies from latitudinal and elevational range limits: a potential adaptation to solar irradiance

1. This study provides evidence that a heliophilic butterfly, the Glanville fritillary (Melitaea cinxia) has adapted differently to environmental variation across latitudes and elevations. 2. In cool air, basking M. cinxia orient themselves perpendicular to the sun's rays to gain heat and take off. During flight, solar heating is reduced because orientation perpendicular to the sun is no longer possible and convective cooling occurs. Consequently, M. cinxia have been shown to suffer net heat loss in flight, even in full sunshine. When flight duration is restricted in this way, the takeoff temperature becomes an important thermal adaptation. 3. Using a thermal imaging camera, takeoff temperatures were measured in experimental butterflies. Butterflies from the northern range limit in Finland took flight at slightly hotter temperatures than butterflies from the southern limit in Spain, and much hotter than butterflies from the elevational limit (1900–2300 m) in the French Alps. Butterflies from low‐elevation populations in southern France also took off much hotter than did the nearby Alpine population. 4. These results suggest that the influence of elevation is different from that of latitude in more respects than ambient temperature. Values of solar irradiance in the butterflies' flight season in each region show that insects from the coolest habitats, Finland and the Alps, experienced similar solar irradiance during basking, but that Finns experienced much lower irradiance in flight. This difference may have favored Finnish butterflies evolving higher takeoff temperatures than Alpine butterflies that also flew in cool air but benefited from more intense radiant energy after takeoff.