Wild herbivores and cattle have differing effects on postfire herbaceous vegetation recovery in an African savanna.

Fire and herbivory have profound effects on vegetation in savanna ecosystems, but little is known about how different herbivore groups influence vegetation dynamics after fire. We assessed the separate and combined effects of herbivory by cattle and wild meso- and megaherbivores on postfire herbaceous vegetation cover, species richness, and species turnover in a savanna ecosystem in central Kenya. We measured these vegetation attributes for five sampling periods (from 2013 to 2017) in prescribed burns and unburned areas located within a series of replicated long-term herbivore exclosures that allow six different combinations of cattle and wild meso- and megaherbivores (elephants and giraffes). Vegetation cover (grasses, mainly) and species richness were initially reduced by burning but recovered by 15-27 months after fire, suggesting strong resilience to infrequent fire. However, the rates of recovery differed in plots accessible by different wild and domestic herbivore guilds. Wildlife (but not cattle) delayed postfire recovery of grasses, and the absence of wildlife (with or without cattle) delayed recovery of forbs. Herbivory by only cattle increased grass species richness in burned relative to unburned areas. Herbivory by cattle (with or without wildlife), however, reduced forb species richness in burned relative to unburned areas. Herbivory by wild ungulates (but not cattle) increased herbaceous species turnover in burned relative to unburned areas. Megaherbivores had negligible modifying effects on these results. This study demonstrates that savanna ecosystems are remarkably resilient to infrequent fires, but postfire grazing by cattle and wild mesoherbivores exerts different effects on recovery trajectories of herbaceous vegetation.

Effects of altering diet carbohydrate profile and physical form on zoo-housed giraffe Giraffa camelopardalis reticulata.

Unlike wild giraffe that primarily consume low starch browse, the preference of zoo-housed giraffe for consuming supplemental feeds over forage could increase the risk of digestive disorders such as ruminal acidosis. Our objective was to evaluate the effects of modifying a supplemental feed's non-fibre carbohydrate profile and physical form on nutritional, behavioural, and blood measures of giraffe in a zoological setting. Six non-lactating, adult, female reticulated giraffes were used in a two-pen modified reversal study using two dietary treatments in seven 21-day periods with data collected on days 15-21. Dietary treatments were a control feed comprised of commercially available products used at the time as the giraffe feed (GF) and an unpelleted experimental feed (EF). On a dry matter basis, GF and EF, respectively, contained 17.0% and 17.4% crude protein, 14.2% and 1.5% starch, 14.9% and 21.3% ethanol-soluble carbohydrates, 22.9% and 26.0% acid detergent fibre (ADF) and 9.50% and 14.9% ND-soluble fibre (NDSF), with modulus of fineness values of 3.62 and 4.82. Supplemental feeds, alfalfa hay, salt, and water were available for ad libitum consumption. Significance was declared at p ≤ 0.05. Intakes of hay, supplemental feeds, and total feed did not differ by diet (p > 0.28), though intakes of starch (0.93 and 0.12 kg; p = 0.05) and ADF (1.83 and 2.23 kg; p = 0.04) differed between GF and EF respectively. Giraffe behaviour values (min/48 h) were greater with EF for total eating (p = 0.04); diets were not detected as different for engagement in oral stereotypes (GF = 433, EF = 318 min/48 h; p = 0.22). Blood glucose was higher on GF than EF (99.0 and 82.3 mg/dL; p = 0.03). The lower EF blood glucose value is more similar to ranges reported for domesticated ruminants. No differences were detected for changes in body weight or body condition score in the 21-day periods (p > 0.32). Modification of supplemental feed carbohydrate profile and physical form can influence behaviour and blood glucose values of zoo-housed giraffe.

Factors affecting captive female giraffe stress response: Male presence, small enclosure, and low temperature.

To improve animal welfare based on suitable social housing conditions, it is important to understand the factors that trigger high-stress responses. Wild giraffes live in a fission-fusion society and males and females are rarely in the same herd for a long period. The captive condition of belonging to a herd with the same individuals for months or years is uncommon in nature. To understand the effect of male presence on female stress levels, fecal glucocorticoid metabolite (fGCM) levels and social interactions in two captive female giraffes were investigated. Additionally, the effect of enclosure size and temperature on fGCM level and social interactions were examined. The results showed no significant difference in the fGCM levels of females based on male presence. The frequency of agonistic behavior by the dominant female toward the subordinate female was significantly increased when a male was present. The subordinate female was significantly less likely to approach the dominant female and showed decreased affiliative and agonistic interactions toward the dominant female when a male was present. The frequencies of agonistic interactions between females were higher in the small enclosure regardless of male presence. Low temperature triggered higher fGCM levels and increased agonistic interaction in an aged female. The findings of this study suggest that these multiple factors should be considered individually to promote the welfare of captive giraffes.

Response to comment on "Sexual selection promotes giraffoid head-neck evolution and ecological adaptation".

Hou et al. challenged the giraffoid affinity of Discokeryx and its ecology and behavior. In our response we reiterate that Discokeryx is a giraffoid that, along with Giraffa, shows extreme evolution of head-neck morphologies that were presumably shaped by selective pressure from sexual competition and marginal environments.

Comment on "Sexual selection promotes giraffoid head-neck evolution and ecological adaptation".

Wang et al. (Research Articles, 3 June 2022, eabl8316) reported an early Miocene giraffoid that exhibited fierce head-butting behavior and concluded that sexual selection promoted head-neck evolution in giraffoids. However, we argue that this ruminant is not a giraffoid and thus that the hypothesis that sexual selection promoted giraffoid head-neck evolution is not sufficiently supported.

The cardiovascular challenges in giraffes.

Giraffes are the highest living animals on Earth and therefore are challenged by gravity more than any other species. In particular the cardiovascular system needs to adapt to this challenge. Giraffes have a mean blood pressure around 200 mmHg, which ensures a mean arterial pressure near the head of 100 mmHg when the giraffe is standing with the neck in a near vertical position. This immediately raises several questions. How do giraffes avoid edema in the legs where the arterial pressure is 300 mmHg or higher? How does the heart produce a pressure of 200 mmHg, and what is the energy required for this endeavor? How can the kidney tolerate a pressure of about 200 mmHg and does this mean that giraffes have a high glomerular filtration rate? What is the arterial pressure in the head of giraffes when they drink, and how is perfusion of the brain maintained when they lift their head after drinking? In this short review, we present some answers to these questions.

MANDIBULAR FRACTURES IN GIRAFFES (GIRAFFA CAMELOPARDALIS) IN EUROPEAN ZOOS.

Following a case of mandibular fracture in a giraffe (Giraffa camelopardalis) at the Budapest Zoo, a survey was conducted across European zoological institutions with the support of the European Association of Zoos and Aquariums Giraffe Ex situ Programme (EAZA Giraffe EEP) to help identify the incidence, potential causes, and management of such injuries. Two hundred sixteen zoos keeping giraffes were invited to participate in the survey. Eighty-six responses were received, 14 of which (16.3%) reported jaw fractures with various causes in their giraffes. Of these responses, hay racks, where the muzzle of the giraffe could fit between the grid of feeding contraptions, were associated with mandibular fractures in seven cases (50.0%). Most giraffes were managed surgically (10/14; 71.4%), two were managed medically (14.3%), one was euthanized because of the severity of the injury (7.1%), and one was only diagnosed with a mandibular fracture postmortem. One giraffe died owing to postanesthetic complications; all other individuals had a full recovery. Most mandible fractures occurred in relatively young giraffes (between 5 wk and 8 yr with an average of 3.4 yr). Based on these findings, facilities with giraffe should evaluate their hay feeders for mandible entrapment risk, especially if they house young individuals.

Effects of browsing enrichment associated with the temperature-humidity index and landscaping trees in giraffes (Giraffa camelopardalisreticulata).

Enclosure environments for captive giraffes can be improved by promoting species-specific behaviors and extending foraging behavior. To date, however, the effects of climatic (temperature-humidity index, THI) and environmental factors (landscaping trees) on the enrichment of captive settings have not been studied. Therefore, the present study explored the effects of browsing enrichment on the licking behavior of captive giraffes. From May 2019 to February 2020, three giraffes in the Kyoto City Zoo, Japan, were observed for 270 h over two consecutive seasons (pre- and post-deciduous). Overall, branch foraging behavior and licking behavior were weakly and negatively correlated. In the pre-deciduous period, THI was significantly and negatively correlated with giraffe activity, and some individuals exhibited significantly reduced foraging behavior. Therefore, browsing enrichment under extreme heat may not improve giraffe rearing environments. Moreover, in the post-deciduous period, with decreased availability of leaves on landscaping trees, the non-branch foraging behavior of giraffes was significantly increased, with a corresponding significant increase in licking behavior. Therefore, landscaping trees affect the foraging behavior of captive giraffes.

Further behavioural parameters support reciprocity and milk theft as explanations for giraffe allonursing.

Suckling of a non-filial calf, or allosuckling, is an extreme case of allomaternal care in mammals. There have been many hypotheses postulated in an attempt to explain this behaviour; however, the supporting evidence differs, together with the conclusions drawn from the investigated variables. Previously, suckling bout rejection was analysed, and the milk theft and reciprocity hypotheses were both determined as the most appropriate explanation of allosuckling in giraffe. In this study, seven hypotheses were tested using different behavioural parameters, namely suckling bout frequency, suckling bout duration, and time spent suckling. It is well-documented that these parameters are associated with various aspects in ungulate biology; for example, suckling rejection typically reflects milk intake and parent-offspring conflict, whereas the suckling bout duration and frequency is associated with social behaviours (affiliation, bonding, social stress). In total, 22 nursing females and 47 suckling calves were observed, in four Czech zoos during a five-year period. The correlation of the observed parameters between the reciprocal female-calf dyads was found to be in line with the reciprocity hypothesis. In addition, non-filial calves tried to steal the milk from non-maternal females, supporting the milk theft hypothesis. Thus, the results support both the reciprocity and milk-theft hypotheses as the most plausible explanation of allosuckling in giraffe, and illustrates the importance of using suckling bout duration and frequency, and the time spent suckling, as behavioural parameters that may aid in explaining the extremity of maternal investment, such as allosuckling.

Biological and environmental factors as sources of variation in nocturnal behavior of giraffe.

Upon a drastic decline of the giraffe population in the wild, conservation efforts and therefore the role of zoos have become more important than ever. With their unique opportunities, zoos provide excellent conditions to study animal behavior, expanding the knowledge about the giraffe's behavior repertoire and their ability to adapt. This study therefore examined the nocturnal behavior of 63 giraffe living in 13 different EAZA zoos across Germany and the Netherlands. Giraffe were observed and videos recorded via infrared sensitive cameras during the winter seasons 2015-2018. The observation period spanned nightly from 17:00 to 7:00. Thus, 198 nights, with a total of 2772 h were recorded and analyzed. Linear mixed models were then used to assess potential biological and environmental factors influencing behavior during the dark phase. Results show that individual variables such as age, subspecies and motherhood determined nocturnal activity and sleep behavior most. Among the variables studied, husbandry conditions and environmental factors complying with EAZA standards had no influence on the giraffe's nocturnal behavior. By combining nocturnal activity analyses and an assessment of potential influencing factors, our findings present a holistic approach to a better understanding of captive giraffe behavior and allow for management implications.

Giraffes and hominins: reductionist model predictions of compressive loads at the spine base for erect exponents of the animal kingdom.

In humans, compressive stress on intervertebral discs is commonly deployed as a measurand for assessing the loads that act within the spine. Examining this physical quantity is crucially beneficial: the intradiscal pressure can be directly measured in vivo in humans, and is immediately related to compressive stress. Hence, measured intradiscal pressure data are very useful for validating such biomechanical animal models that have the spine incorporated, and can, thus, compute compressive stress values. Here, we use human intradiscal pressure data to verify the predictions of a reductionist spine model, which has in fact only one joint degree of freedom. We calculate the pulling force of one lumped anatomical structure that acts past this (intervertebral) joint at the base of the spine, lumbar in hominins, cervical in giraffes, to compensate the torque that is induced by the weight of all masses located cranially to the base. Given morphometric estimates of the human and australopith trunks, respectively, and the giraffe's neck, as well as the respective structures' lever arms and disc areas, we predict, for all three species, the compressive stress on the intervertebral disc at the spine base, while systematically varying the angular orientation of the species' spinal columns with respect to gravity. The comparison between these species demonstrates that hominin everyday compressive disc stresses are lower than those in big quadrupedal animals. Within each species, erecting the spine from being bent forward by, for example, thirty degrees to fully upright posture reduces the compressive disc stress roughly to a third. We conclude that erecting the spine immediately allows the carrying of extra loads of the order of body weight, and yet the compressive disc stress is lower than in a moderately forward-bent posture with no extra load.

The Remarkable Cardiovascular System of Giraffes.

Gravity affects the physiology of many animals, and the effect is, for good reason, most pronounced in tall species. The physiology-in particular, cardiovascular function-of giraffes has therefore captivated the interest of physiologists for centuries. Several studies document high mean arterial blood pressure of giraffes of about 200 mm Hg. This appears necessary to establish a cerebral perfusion pressure on the order of 100 mm Hg at the cranial end of the carotid arteries. Here, we discuss the unique characteristics of blood vessels, the heart, and the kidney of giraffes and how these functional and structural adaptations are related to very high blood pressure. We also discuss how the cerebral circulation of giraffes is established and what we know about how the blood flow and arterial and venous pressures in giraffes change when they stop to drink and subsequently lift their heads 5-6 m in one sweeping movement.

The impact of Masai giraffe nursery groups on the development of social associations among females and young individuals.

Fission-fusion social systems involve the splitting and merging of subgroups with frequent changes in membership occurring as a result of a number of ecological and social factors, such as demographic processes including birth, movement, or death. Giraffe reside in fission-fusion social systems, and we studied how reproductive status influence associations among females, as well as how associations differ between calves and juveniles. Data were collected in Katavi National Park, Tanzania, during five study periods. We used social network analysis to identify whether reproductive status and developmental stages predict differences in giraffe social association. We found that females with offspring maintain stronger associations than females without offspring. We also revealed that calves and juveniles had similar network association patterns. Our results suggest that the presence of dependent offspring influences the social associations of females and individuals less than 1.5 years of age are still maintaining strong social associations with nursery group members. We conclude that nursery groups among giraffe are co-operative rearing units that probably reduce the costs of rearing to mothers, and may provide a group structure for animals to begin to develop skills useful for their future life in a fission-fusion social system.

Nighttime Suckling Behavior in Captive Giraffe (Giraffa camelopardalis reticulata).

Suckling behaviors are useful to better understand mother-offspring relationships. However, in many species, knowledge about nighttime suckling behavior is sparse. In the present study, we investigated suckling behavior in four calves of the reticulated giraffe (Giraffa camelopardalis reticulata) and their mother in the Kyoto City Zoo, Japan, by analyzing video clips of 9614 h. The relation between a calf's age and the mean duration and frequency of suckling were consistent with the results of previous daytime studies: the longest duration (80 ± 4 s) and highest frequency (0.41/h) of suckling occurred when the calves were young, and decreased thereafter. Suckling frequency was lower at night than during the day, probably because the mother spent much of the time resting at night. The mother initiated suckling behavior in offspring until six months of age. Most often, the mother terminated suckling except during the first few weeks after birth. Additionally, we reported that the average weaning period was 448 days (n = 5) and that the last born (sixth) continued to suckle for more than 750 days, which is a rare phenomenon. We believe the present report helps to fill the lack of knowledge about nighttime suckling behavior in giraffe.

Evaluating the social structure of captive Rothschild's giraffes (Giraffa camelopardalis rothschildi): Relevance to animal management and animal welfare.

Social network analysis (SNA) is useful for evaluating management zoo regimes to ensure that any fitness benefits of sociality are preserved in captive-housed groups. This paper explores the association patterns of 13 giraffes housed at Longleat Safari Park, UK. Wild giraffes exhibit a fission-fusion social system with preferential bonding. As zoo-housed giraffes are common, they are excellent study subjects for using SNA to investigate key aspects of sociality within a managed social environment. Social bonds were assessed over different seasons and data from two study periods (2011 and 2015) were analyzed to see consistency of "social type" (i.e., more social or more solitary). Associations showed the occurrence of consistent preferential bonds between named individuals but time of year influenced the patterns of social bonds. Female-female bonds and female-offspring bonds appeared to be strongest. For animals present in 2011 and 2015, differences in time spent socializing between years were apparent. Results suggest that giraffes may be flexible in their choice of social partner and zoo-managed herds should include a range of individuals from which each animal can choose a preferred associate.

GROWTH, HUSBANDRY, AND DIETS OF FIVE SUCCESSFULLY HAND-REARED ORPHANED GIRAFFE CALVES (GIRAFFA CAMELOPARDALIS ROTHSCHILDI AND GIRAFFA CAMELOPARDALIS RETICULATA).

Giraffe in the wild are in ongoing decline because of poaching and habitat loss and fragmentation, and were recently assessed as "vulnerable" on the IUCN (International Union for Conservation of Nature) Red List of Threatened Species. Captive breeding and saving each individual are therefore becoming more important to save this species from extinction. This paper describes the husbandry and diets of successfully hand-reared Rothschild's giraffes (Giraffa camelopardalis rothschildi; n = 3) and reticulated giraffes (Giraffa camelopardalis reticulata; n = 2). All calves were initially fed with bovine colostrum followed by cow's milk (Holstein milk; Holstein milk with 10% of bovine colostrum; Jersey and Guernsey milk). Additionally, lactase enzymes (Lactaidt, Johnson & Johnson Inc., Guelp, Ontario N1K1A5, Canada) and probiotics (Probiost, Vets Plus, Inc., Menomonie, WI 54751, USA) were used. Average growth varied from 764 to 1,239 g/day from birth until 2 mo of age and between 508 and 1,161 g/day from birth until last measurement before weaning. Hand-reared calves gained up to 21 cm in height within the first month and 82-138% of their birth weight during the first 2 mo. The giraffes were weaned at 6 (n = 1), 8 (n = 3), and 11.5 (n = 1) mo and successfully socialized and introduced to other giraffes. The described diets and husbandry proved to be effective in all five calves. Large amounts of cow's milk per feeding (up to 6 L) did not result in gastrointestinal problems.

Comparisons of Schansitherium tafeli with Samotherium boissieri (Giraffidae, Mammalia) from the Late Miocene of Gansu Province, China.

We are describing and figuring for the first time skulls of Schansitherium tafeli, which are abundant in the Gansu area of China from the Late Miocene. They were animals about the size of Samotherium with shorter necks that had two pairs of ossicones that merge at the base, which is unlike Samotherium. The anterior ossicones consist of anterior lineations, which may represent growth lines. They were likely mixed feeders similar to Samotherium. Schansitherium is tentatively placed in a very close position to Samotherium. Samotherium and Schansitherium represent a pair of morphologically very similar species that likely coexisted similarly to pairs of modern species, where the main difference is in the ossicones. Pairs of ruminants in Africa, for example, exist today that differ mostly in their horn shape but otherwise are similar in size, shape, and diet. The absence of Schansitherium from Europe is interesting, however, as Samotherium is found in both locations. While is it challenging to interpret neck length and ossicone shape in terms of function in combat, we offer our hypothesis as to how the two species differed in their fighting techniques.

The locomotor kinematics and ground reaction forces of walking giraffes.

Giraffes (Giraffa camelopardalis) possess specialised anatomy. Their disproportionately elongate limbs and neck confer recognised feeding advantages, but little is known about how their morphology affects locomotor function. In this study, we examined the stride parameters and ground reaction forces from three adult giraffes in a zoological park, across a range of walking speeds. The patterns of GRFs during walking indicate that giraffes, similar to other mammalian quadrupeds, maintain a forelimb-biased weight distribution. The angular excursion of the neck has functional links with locomotor dynamics in giraffes, and was exaggerated at faster speeds. The horizontal accelerations of the neck and trunk were out of phase compared with the vertical accelerations, which were intermediate between in and out of phase. Despite possessing specialised morphology, giraffes' stride parameters were broadly predicted from dynamic similarity, facilitating the use of other quadrupedal locomotion models to generate testable hypotheses in giraffes.