Pygmy Hippopotamus Research Articles

INDUCTION OF PARTURITION IN A PYGMY HIPPOPOTAMUS (CHOEROPSIS LIBERIENSIS).

A 27-yr-old female pygmy hippopotamus (Choeropsis liberiensis) had two consecutive stillbirths with no overt signs of labor, suggestive of uterine inertia. After a third pregnancy was confirmed, an induction protocol was developed. Cloprostenol and betamethasone were administered on d 200 of gestation (time 0 h). Additional doses of cloprostenol were administered at 24 and 48 h and oxytocin at 30, 31, and 48 h. Each injection resulted in preparturient behavior without overt evidence of contractions. Fetal membranes presented at the vulva at 54.5 h after initial cloprostenol and betamethasone administration with no progression of labor. Transvaginal palpation and manual delivery of a live calf followed. Despite confirmed nursing, the serum glutaraldehyde coagulation test was negative. Failure of passive transfer may have been secondary to the induction protocol. The calf was treated with broad-spectrum antimicrobial agents due to diarrhea, and clinical signs resolved. This clinical brief details the first known induction of parturition in a pygmy hippopotamus, which can serve as the basis for further development of the technique.

Forest loss during 2000–2019 in pygmy hippopotamus (Choeropsis liberiensis) habitats was driven by shifting agriculture

SummaryThe Upper Guinea Forest (UGF; West Africa), a global biodiversity hotspot, has lost more than 90% of its original area since 1900, threatening endemic species such as the endangered pygmy hippopotamus (Choeropsis liberiensis). However, little is known about the proximate causes of this deforestation. We classified Sentinel-2 data using the random forest algorithm to differentiate between three main human processes (shifting agriculture, intensive agriculture or urban expansion) driving deforestation between 2000 and 2019 across the pygmy hippopotamus distribution area. Out of c. 89 600 km2 in the year 2000, 15 900 km2 (17%) of forest were lost, primarily to shifting agriculture (14 900 km2). Côte d’Ivoire and Liberia accounted for 14 900 km2 (94%) of the net area of forest lost, c. 15 times greater than deforestation in Sierra Leone and Guinea combined (953 km2). Forest loss inside protected areas is pervasive, and it is essential to prioritize conservation efforts in areas where deforestation is still low (e.g., Taï, Sapo and Gola Rainforest national parks). We suggest that the preservation of the UGF will face challenges associated with people’s demand for food and income. Continued landscape-scale planning and action to reduce deforestation are urgently needed to limit the impact of shifting agriculture on pygmy hippopotamus habitat.

Chewing, dentition and tooth wear in Hippopotamidae (Hippopotamus amphibius and Choeropsis liberiensis).

Among mammals, hippopotamids ('hippos') have been described as the species with the lowest chewing efficacy despite elaborate enamel folds on the occlusal surface or their cheek teeth, which was hypothesized to result from the lack of a grinding chewing motion. We investigated the chewing and dentition of the two extant hippo species, the common hippo (Hippopotamus amphibius) and the pygmy hippo (Choeropsis liberiensis), making (video) observations of live animals and gathering data on museum specimens (n = 86 H. amphibius and 26 C. liberiensis skulls). Hippos have a low degree of anisodonty (differences in width between maxillary and mandibular cheek teeth) and anisognathy (difference in width between the upper and the lower jaw), corresponding to a mainly orthal (up-and-down) chewing motion. The two hippo species differ slightly, but distinctively, in their anterior dental morphology and chewing mode. In both species, the canines do not completely prevent a lateral jaw movement but would, in theory, permit this movement until the mandibular canines get into contact with the maxillary protruding snout. This movement is only realized, to a small extent, in pygmy hippos, leaving distinct wear traces on their incisors and creating relatively wider wear facets on the maxillary canines. In common hippos, the interlocking upper and lower incisors prevent lateral jaw movement. Corresponding contact wear facets are evident on the medial aspect of the upper, and on the lateral aspect of the lower incisors-unless museal reconstructions mispositioned these teeth. If these facets are interpreted as an indication for a relic of a lateral jaw movement that was probably more prominent in hippo ancestors, i.e. if we assume that hippos evolved orthal chewing secondarily, several other characteristics of hippos can be explained, such as a low degree of hypsodonty (in the absence of distinct attrition due to a grinding chewing movement), a secondary loss of complexity in their enamel schmelzmuster, a secondary evolution of a wide mouth gape, a reduction in anisodonty compared to their ancestors, and the evolution of a bilaterally symmetrical ('trifoliate') enamel folding pattern on the molar occlusal surface from an ancestral bunoselenodont condition. As an underlying driving force, selection for intraspecific combat with canines and incisors, necessitating a wide gape and a rigid jaw, has been suggested.

Breast Milk Oligosaccharides Contain Immunomodulatory Glucuronic Acid and LacdiNAc

Breast milk is abundant with functionalized milk oligosaccharides (MOs) to nourish and protect the neonate. Yet we lack a comprehensive understanding of the repertoire and evolution of MOs across Mammalia. We report ∼400 MO-species associations (>100 novel structures) from milk glycomics of nine mostly understudied species: alpaca, beluga whale, black rhinoceros, bottlenose dolphin, impala, L'Hoest's monkey, pygmy hippopotamus, domestic sheep, and striped dolphin. This revealed the hitherto unknown existence of the LacdiNAc motif (GalNAcβ1-4GlcNAc) in MOs of all species except alpaca, sheep, and striped dolphin, indicating the widespread occurrence of this potentially antimicrobial motif in MOs. We also characterize glucuronic acid-containing MOs in the milk of impala, dolphins, sheep, and rhinoceros, previously only reported in cows. We demonstrate that these GlcA-MOs exhibit potent immunomodulatory effects. Our study extends the number of known MOs by >15%. Combined with >1900 curated MO-species associations, we characterize MO motif distributions, presenting an exhaustive overview of MO biodiversity.

Perioperative management of umbilical herniorrhaphy with ileal resection and anastomosis in a pygmy hippopotamus (Choeropsis liberiensis)

AbstractA 28‐year‐old 278 kg female pygmy hippopotamus (Choeropsis liberiensis) with a chronic, static umbilical hernia presented acutely for lethargy and anorexia. Ultrasonography and abdominocentesis were consistent with an incarcerated umbilical hernia. An exploratory laparotomy, ileal resection and anastomosis, and herniorrhaphy were carried out via a ventral midline incision and the hippopotamus remained hospitalised for 9 weeks postoperatively. A second surgery was necessary to debride the incision site after the primary skin closure dehisced. Improved support to skin sutures was provided using sterilised plastic cable zip ties, which were better suited to the unique skin of the hippopotamus and tension associated with the animal's size and anatomy. Incisional closure is a recognised challenge in this species and significant reinforcement is required to prevent wound dehiscence. The novel use of cable zip ties provided stability where suture material had transected the incision and should be considered in hippopotamus surgery.

What can hippopotamus isotopes tell us about past distributions of C4 grassy biomes on Madagascar?

Societal Impact StatementToday, expansive C4 grassy biomes exist across central, western, and northern Madagascar. Some researchers have argued that the island's now‐extinct pygmy hippopotamuses belonged to a megaherbivore grazing guild that maintained these grasslands prior to human arrival. However, the chemistry of hippo bones indicates that C4 grasses were only a minor part of hippo diet. This, in turn, suggests that C4 grasses were present but not widespread when hippos were alive and that grasses expanded only after Malagasy people shifted from hunting and foraging to agropastoralism approximately 1000 years ago. These results have important implications for environmental reconstructions and biodiversity management.Summary Extinct hippopotamuses (Hippopotamus spp.) were part of Madagascar's megaherbivore guild. Stable carbon (δ13C) and nitrogen (δ15N) isotopes in radiometrically dated bone collagen track spatial and temporal variation in diet and habitat. If hippos helped maintain C4 grassy biomes, then they should have regularly consumed C4 grasses, which have high δ13C values. However, if expansive C4 grassy biomes are anthropogenic, then forests would have been more extensive in the past, and hippos would have predominantly consumed C3 plants with low δ13C values. Nitrogen isotopes can clarify foraging habitat (moist or dry). We assessed δ13C and δ15N values for hippos from different ecoregions of Madagascar and compared these with data for extinct herbivorous lemurs from the same ecoregions. We further explored the effects of wet/dry transitions on isotopic trends for hippos from the central highlands and spiny thicket ecoregions. Carbon isotopes suggest (1) limited C4 consumption by hippos in the central highlands, dry deciduous forest, and succulent woodland ecoregions; and (2) moderate consumption of C4 resources in the spiny thicket. Nitrogen data indicate that hippos foraged in wetter habitats than sympatric lemurs in all regions. Malagasy hippos did not regularly graze C4 grasses in dry, open habitats, even in regions blanketed by C4 grassy biomes today. Malagasy grasses are adapted to grazing and fire, but these are likely ancient adaptations that accompanied grasses when they initially spread to Madagascar. C4 grassy biomes were spatially limited in extent in the past and only expanded after the Late Holocene introduction of domesticated ungulates.

Molecular data suggests a Pliocene – Early Pleistocene divergence date for the common and pygmy hippopotamus

Hippopotamid phylogeny has proven difficult to resolve with proposed relationships between extant and fossil species receiving mixed levels of support. Of particular interest is the divergence between the two extant hippopotamid species, the well-known common hippopotamus (Hippopotamus amphibius) and the enigmatic pygmy hippopotamus (Choeropsis liberiensis). Previous studies have relied on morphological and fossil evidence to identify the ancestral species at the core of this divergence and its timing. In this study, we assembled a molecular matrix of 26 nuclear gene sequences from 11 ungulate species with two primates as an outgroup. We used a Bayesian relaxed molecular clock approach to reconstruct a calibrated time tree for Cetartiodactyla and estimate the divergence date between the common and pygmy hippopotamus. While previous morphological studies have estimated this event to have occurred sometime during the Late Miocene (between 11.6 and 5.3 million years ago), our nuclear gene-based estimates suggest a more recent split of about 4.04 Ma (95% confidence interval: 8.31–1.97 Ma) via RelTime-ML or 2.4 Ma (95% confidence interval: 3.1–1.6 Ma) via MCMCTree. These more recent estimates correspond with the Early Pliocene – Early Pleistocene sub-epochs and align most closely with the results of previous genomic studies. We discuss how our results compare with previous estimates based on both morphological and molecular studies, some of which extend the predicted range of this divergence date even further back in time. Our results suggest a different path of evolution for the understudied pygmy hippopotamus and reveal that morphological evidence alone may not resolve the correct hippopotamid phylogenetic and time trees. We suggest that the common and pygmy hippopotamus may be phylogenetically closer than once believed. Our results also call for further studies to develop a combined approach incorporating both molecular and morphological evidence to reach a consensus on the evolutionary patterns and timing that led to modern hippopotamid evolution.

Non-invasive assessment of fecal glucocorticoid and androgen metabolites in the pygmy hippopotamus (Choeropsis liberiensis)

The pygmy hippopotamus (Choeropsis liberiensis) is an endangered species endemic to the Upper Guinea Forest ecosystem in West Africa. We have limited information concerning the species’ reproduction and well-being under managed care. We therefore developed non-invasive methods for characterizing gonadal androgen and adrenal hormone profiles in pygmy hippos using fecal samples collected from 12 males and 12 females housed in North American zoological institutions. We aimed to: 1) identify and validate enzyme immunoassays (EIAs) for measuring metabolites of corticosteroids and testosterone in feces; and 2) test whether gonadal activity is correlated with previous breeding history, season or type of housing. For glucocorticoids, several EIAs for measuring metabolites were investigated. A group-specific EIA exhibiting cross-reactivity with 11,17-dioxoandrostane (DOA) metabolites of cortisol most clearly reflected adrenocortical activity in response to pharmocological challenge with adrenocorticotropic hormone (ACTH) in both males and females. However, day-to-day concentrations of this metabolite in the feces of pygmy hippos that did not undergo ACTH challenge were near the detection limits of the assay, making this EIA impractical for assessing glucocorticoid activity in this species. Another group-specific EIA, exhibiting cross-reactivity with 5α-pregnane-3β,11β,21-triol-20-one, produced biologically relevant data and evidence of an appropriate response to pharmacological challenge with exogenous ACTH. The testosterone metabolite assay C196 (Arbor Assays, Ann Arbor, Michigan, USA) also produced biologically coherent data: adult males exhibited the highest mean androgen metabolite concentrations (477 ng/g), followed by adult females (259 ng/g) and juvenile males (160 ng/g). Proven breeding males had higher, but not significantly different, mean concentrations (472 ng/g) to unproven males (352 ng/g; P = 0.400). Similarly, adult males housed outdoors year-round in subtropical climates exhibited higher, but not statistically different mean concentrations (554 ng/g) to males in temperate climates that were housed indoors at least part of the year (412 ng/g; P = 0.208). There were, however, significant differences in mean concentrations among seasons for adult males, with higher values in spring (546 ng/g) and summer (542 ng/g) than in autumn (426 ng/g) and winter (388 ng/g, P = 0.003). In conclusion, we identified EIAs for the measurement of fecal metabolites of androgens and glucocorticoids that can be used for further studies to monitor gonadal activity in male pygmy hippos and adrenocortical activity in both sexes. We also identified a seasonal trend in male gonadal activity in this species under managed care in North America. Finally, our findings highlight an important consideration when using non-invasive methods for evaluating fecal cortisol metabolites: ACTH used for pharmacological validation of an EIA does not necessarily equate to biological relevance.

From hippos to cattle: The fragile relationship of early cypriots with their animals

The relationship between humans and animals has always been complex. Two seminal topics researchers have examined in detail relate to human-induced extinctions and to the major change in human/animal interactions during the domestication of certain key species. This essay examines animals and the earliest humans on the Mediterranean island of Cyprus from two perspectives. The first relates to the late Epipaleolithic where humans likely were instrumental in the extinction of endemic species, primarily the pygmy hippopotamus. The second focuses on the early Neolithic, when animals were coming under domestication. During this time, people still relied heavily on wild species (deer) and may have used some domesticates in a feral sense. These complex relationships demonstrate that Cyprus, despite its relatively small land mass, was involved in the sophisticated utilization of animals since the first humans were on the island.

Comparison of the Rostral Epidural Rete Mirabile and the Patterns of Its Blood Supply in Selected Suiformes and Hippopotamuses.

The rostral epidural rete mirabile (rete mirabile epidurale rostrale) is built of considerable small arterial vessels that anastomose with each other. This structure is formed as a result of the division of a large arterial vessel into many small arteries. Furthermore, on the other side of the rete mirabile, these small arteries are joined together to form one large artery, through which blood flows out of the rete mirabile. This system participates in decreasing the temperature of the brain and thus protects the body from thermal stress. Moreover, it influences body-water balance and has a crucial role in the retrograde transfer of neuropeptides. The goal of this study was to describe the rostral epidural rete mirabile and pathways that provide blood to this structure as well as compare it in selected Suiformes and hippopotamuses. The study was performed on desert warthogs (Phacochoerus aethiopicus), Eurasian wild boars (Sus scrofa), collared peccaries (Pecari tajacu), pygmy hippopotamuses (Choeropsis liberiensis), and common hippopotamuses (Hippopotamus amphibius). Preparations were made using the latex method and corrosion cast. An elongated shape characterizes its anatomy with a much wider rostral part than caudal part in the Eurasian wild boars, desert warthogs, and collared peccaries. The main source of blood was the branch to the rostral epidural rete mirabile branched off from the internal carotid artery. Moreover, blood enters the rete by the caudal branch and rostral branch to the rostral epidural rete mirabile. In hippopotamuses, the major source of blood was the rostral branches to the rostral epidural rete mirabile.

Oxygen isotope analysis of pygmy hippo’s fossil bone and tooth apatite from Aghia Napa, Cyprus

Dwarf elephants, pygmy hippos and various species of micromammals are some of the animals that lived on the island of Cyprus, 11,000 to 13,000 years ago. Fossil skeletal material of the pygmy hippopotamus Phanourios minor has been excavated from various fossiliferous sites in Cyprus, including Aghia Napa. Phanourios minor is considered to be the smallest of all known insular hippopotamuses. The animal is estimated to have measured about 76 cm (2.5 ft) tall and 121 cm (4.0 ft) long. During our study, material from Aghia Napa was used in order to attempt to reconstruct the palaeoclimatic conditions at the time this species lived on the island. The method used to determine the living conditions of this extinct species is the isotopic analysis of the bioapatite of bone and tooth samples.According to our results, and in comparison, to data from other similar studies, the average temperature in Cyprus was probably lower than 24 °C and in general the palaeoclimate (based on temperature) in Aghia Napa, Cyprus 12,000 years ago, was not very different to what it is currently, even though the age of the site coincides with the Younger dryas cold event. This has also been reported by other isotopic studies from Greece and the SE Mediterranean region for the specific era, according to which a cold climate is indicated, slowly passing on to a warmer period. According to today's data, we are now experiencing a warm period. The question that remains is how and to which extent, the SE Mediterranean region was actually affected by the cold event.

Cutmarked bone of drought-tolerant extinct megafauna deposited with traces of fire, human foraging, and introduced animals in SW Madagascar

People could have hunted Madagascar’s megafauna to extinction, particularly when introduced taxa and drought exacerbated the effects of predation. However, such explanations are difficult to test due to the scarcity of individual sites with unambiguous traces of humans, introduced taxa, and endemic megaherbivores. We excavated three coastal ponds in arid SW Madagascar and present a unique combination of traces of human activity (modified pygmy hippo bone, processed estuarine shell and fish bone, and charcoal), along with bones of extinct megafauna (giant tortoises, pygmy hippos, and elephant birds), extirpated fauna (e.g., crocodiles), and introduced vertebrates (e.g., zebu cattle). The disappearance of megafauna from the study sites at ~ 1000 years ago followed a relatively arid interval and closely coincides with increasingly frequent traces of human foraging, fire, and pastoralism. Our analyses fail to document drought-associated extirpation or multiple millennia of megafauna hunting and suggest that a late combination of hunting, forest clearance, and pastoralism drove extirpations.

Functional morphology of the hindlimb of fossilized pygmy hippopotamus from Ayia Napa (Cyprus).

We studied the functional morphology of the postcranial skeleton of the endemic hippopotamus Phanourios minor, derived from the Upper Pleistocene site of Ayia Napa. The deposit, which consists of a hard limestone substrate on which the species moved, has yielded a great abundance of hippopotamus material, making the Ayia Napa locality one of the most important paleontological sites in Cyprus. The immigration of the large-sized mainland Hippopotamus to Cyprus led to the emergence of a new insular species with its main characteristic being the extremely reduced body size. In this study, all the hindlimb elements of the Cypriot hippo are described in detail and compared with those of the modern species, with the extant Hippopotamus amphibius being considered similar to the possible ancestor of P. minor. In some cases, the morphological comparison is reinforced using bones of other extinct insular and mainland hippos. Additionally, we provided a functional analysis of the hindlimb joints, suggesting specific locomotor habits for the species. The anatomical examination reveals that the elements in P. minor are robust with marked muscular insertion areas resembling those found in Hippopotamus. However, there are also similarities with Choeropsis liberiensis in certain morphofunctional traits. P. minor adapted to slow but powerful locomotion with remarkable stabilization, particularly in the zeugopodium and the autopodium. The knee was less mobile in the craniocaudal direction compared with that in recent hippos, while the abduction-adduction movements of the thigh were advanced. The pes presented good mobility in the sagittal plane and limitation in transversal movements. Thus, P. minor displayed modifications to its limbs, influenced by the mountainous island environment and the body size reduction, resulting in specialized locomotion, which was different from that of extant hippopotamuses.

Animal reactivity to camera traps and its effects on abundance estimate using distance sampling in the Taï National Park, Côte d'Ivoire.

The use of camera traps (CTs) has become an increasingly popular method of studying wildlife, as CTs are able to detect rare, nocturnal, and elusive species in remote and difficult-to-access areas. It thus makes them suited to estimate animal density and abundance, identify activity patterns and new behaviours of animals. However, animals can react when they see the CTs and this can lead to bias in the animal population estimates. While CTs may provide many advantages, an improved understanding of their impacts on individual’s behaviour is necessary to avoid erroneous density estimates. Yet, the impact of CTs on detected individuals, such as human odour near the device and the environment, or the infrared illumination, has received relatively little attention. To date, there is no clear procedure to remove this potential bias. Here, we use camera trap distance sampling (CTDS) to (1) quantify the bias resulting from the different animal responses to the CTs when determining animal density and abundance, and (2) test if olfactory, visual and auditory signals have an influence on the animals’ reaction to CTs. Between March 2019 and March 2020, we deployed CTs at 267 locations distributed systematically over the entire Taï National Park. We obtained 58,947 videos from which we analysed four medium- to-large-bodied species (Maxwell’s duiker (Philantomba maxwellii), Jentink’s duiker (Cephalophus jentinki), pygmy hippopotamus (Choeropsis liberiensis) and Western chimpanzee (Pan troglodytes verus)) displaying different behaviours towards the CTs. We then established species-specific ethograms describing the behavioural responses to the CTs. Using these species-specific responses, we observed that the Maxwell’s duiker reacted weakly to CTs (about 0.11% of the distance data), contrary to Jentink’s duiker, pygmy hippopotamus and Western chimpanzee which reacted with relatively high frequencies, representing 32.82%, 52.96% and 16.14% of the distance data, respectively. Not taking into account the species-specific responses to the CTs can lead to an artificial doubling or tripling of the populations’ sizes. All species reacted more to the CTs at close distances. Besides, the Jentink’s duiker and the pygmy hippopotamus reacted significantly more to the CTs at night than during the day. Finally, as for olfactory signals, the probability of reaction to the CTs during the first days after CTs installation was weak in Maxwell’s duiker, but concerned 18% of the video captures in Western chimpanzees which decreasing with time, but they remained high in pygmy hippopotamus and Jentink’s duiker (65% and 70% of the video captures respectively). Careful consideration should be given to animal’s response to CTs during the analysis and in the field, by reducing human’s impact around the CTs installation.

Medium‐to‐large mammal diversity and status of forest antelopes in the Sapo National Park, Liberia

AbstractSapo National Park, Liberia's largest protected area, comprises an area of 1804 km2 within the Upper Guinea forest ecosystem, a global biodiversity hot spot. Our baseline camera‐trap study revealed 32 medium‐to‐large mammal species, including ten Upper Guinea forest endemics, within the park. Results show important populations of threatened antelopes, particularly zebra duiker and Jentink's duiker, along with bay duiker, black duiker, Ogilby's duiker and Maxwell's duiker. Water chevrotain and lowland bongo were also recorded. Camera‐trap images of giant and white‐bellied pangolins, western chimpanzee, pygmy hippopotamus and forest elephant collected during this study further confirm the high conservation value of the site. The ecology of the antelope community is investigated, noting the diurnal activity pattern of zebra duiker and the nocturnal activity of Jentink's duiker, in contrast to previous studies. Both species preferred lower altitude primary forest. Ogilby's duiker was the only antelope that showed a preference for higher altitude forest and was found to be more common than the similar‐sized black duiker and bay duiker. Sapo National Park faces many threats including habitat degradation, poaching for bushmeat and animal parts, and illicit gold mining. Commercial logging and agriculture in the buffer and surrounding forested areas threatens connectivity and increases human–wildlife conflict.

The unique rectus extraocular muscles of cetaceans: Homologies and possible functions.

Each rectus extraocular muscle in cetaceans divides into two portions: a massive palpebral belly that inserts into the deep surface of the eyelids and a smaller scleral belly that inserts onto the eyeball. While the cetacean palpebral insertions have long been recognized, their homologies and functions remain unclear. To compare cetacean rectus EOM insertions with the global and orbital rectus EOM insertions of other mammals we dissected orbital contents of 20 odontocete species, 2 mysticete species and 18 non-cetacean species, both aquatic and terrestrial. Four cetacean species were also examined with magnetic resonance imaging (MRI). All four rectus muscles in cetaceans had well-developed palpebral bellies and insertions. Adjacent palpebral bellies showed varying degrees of fusion, from near independence to near complete fusion. Fusion was most complete towards palpebral insertions and less towards origins. A medial moiety of the superior rectus palpebral belly is likely the levator palpebrae superioris. Smaller but still robust scleral insertions were present on all recti, with the medial rectus (MR) being significantly more muscular than the others. All non-cetacean species examined had recti with distinct global and orbital insertions, the latter generally onto Tenon's capsule. Orbital insertions in pygmy hippopotamus and Florida manatee extended into the deep surfaces of the eyelids, hence qualifying as palpebral insertions. Our results suggest that rectus EOMs of mammals generally have both global and orbital insertions, and that palpebral bellies of cetaceans and other species are modified homologs of the orbital insertions. The presence of palpebral insertions in pygmy hippopotamus and absence in other cetartiodactyls suggests an intermediate condition between terrestrial cetartiodactyls and cetaceans. Palpebral insertions in Florida manatee and reports of their presence in some pinnipeds suggest parallel evolution in multiple aquatic lineages. Various functions of cetacean palpebral recti have been proposed, including eyelid dilators, protection during diving and thermogenesis for warming eye and brain. For further insight into their possible functions, we observed eye movements of captive bottlenose dolphins (Tursiops truncatus) at the U.S. National Aquarium. Our observations showed that in addition to rotation of the eyeball the entire surrounding palpebral region also moves during gaze changes. For example during upward gaze the globe not only rotates in supraduction but translates dorsally as well. It appears the rectus palpebral bellies are responsible for flexing the palpebral structures and thus also translating the globe, while the scleral insertions act directly for ocular rotation. Along with frequent non-conjugate eye movements, the oculomotor mechanics and repertoire of cetaceans are thus quite distinctive. Summarily, axial displacement within the orbit is a major 'eye movement' in cetaceans, with protrusion and retraction mediated by well-developed circular muscles and retractor bulbi respectively. Torsional eye movements driven by elaborate oblique EOMs are likewise significant. The roles of rectus EOMs for ocular rotation via their scleral insertions, especially the highly muscular MR, are for typical supra/infraductions and nasal/temporal ductions. The palpebral bellies accentuate these ductions by translating the globe and surrounding structures in the same direction.

Oxygen Isotope Analysis of Pygmy Hippo's Fossil Bone and Tooth Apatite from Aghia Napa, Cyprus

Oxygen Isotope Analysis of Pygmy Hippo's Fossil Bone and Tooth Apatite from Aghia Napa, Cyprus

Mitochondrial sequences of the extinct Cypriot pygmy hippopotamus confirm its phylogenetic placement

Abstract The Cypriot pygmy hippopotamus is an extinct Mediterranean species that inhabited the island of Cyprus during the Late Pleistocene. This iconic species last appears at the archaeological site of Akrotiri Aetokremnos, dated to ~12 500 cal. BP. Taxonomically, the Cypriot pygmy hippopotamus has been assigned to Hippopotamus minor, which, based on morphology, is more closely related to the common hippopotamus (Hippopotamus amphibius; today present only in Sub-Saharan Africa) than to the West African pygmy hippopotamus (Choeropsis liberiensis). Despite adverse conditions for preservation that greatly reduce the likelihood of DNA preservation in subfossil samples from hot environments, we recovered and analysed ancient DNA from petrous bones excavated from Akrotiri Aetokremnos. By whole mitogenome hybridization capture and exhaustive high-throughput sequencing, we were able to investigate the molecular phylogeny and taxonomic status of the Cypriot pygmy hippopotamus. The results of our low-coverage ancient mitogenomic analyses support the close phylogenetic affinity of H. minor to H. amphibius, with their divergence estimated at ~1.36 or 1.58 Mya, depending on the molecular dating method. To our knowledge, this study constitutes the first step towards reconstruction of the molecular phylogeny of Mediterranean Hippopotaminae.

Drought Coincided with, but Does Not Explain, Late Holocene Megafauna Extinctions in SW Madagascar

Climate drying could have transformed ecosystems in southern Madagascar during recent millennia by contributing to the extinction of endemic megafauna. However, the extent of regional aridification during the past 2000 years is poorly known, as are the responses of endemic animals and economically important livestock to drying. We inferred ~1600 years of climate change around Lake Ranobe, SW Madagascar, using oxygen isotope analyses of monospecific freshwater ostracods (Bradleystrandesia cf. fuscata) and elemental analyses of lake core sediment. We inferred past changes in habitat and diet of introduced and extinct endemic megaherbivores using bone collagen stable isotope and 14C datasets (n = 63). Extinct pygmy hippos and multiple giant lemur species disappeared from the vicinity of Ranobe during a dry interval ~1000–700 cal yr BP, but the simultaneous appearance of introduced cattle, high charcoal concentrations, and other evidence of human activity confound inference of drought-driven extirpations. Unlike the endemic megafauna, relatively low collagen stable nitrogen isotope values among cattle suggest they survived dry intervals by exploiting patches of wet habitat. Although megafaunal extirpations coincided with drought in SW Madagascar, coupled data from bone and lake sediments do not support the hypothesis that extinct megafauna populations collapsed solely because of drought. Given that the reliance of livestock on mesic patches will become more important in the face of projected climate drying, we argue that sustainable conservation of spiny forests in SW Madagascar should support local livelihoods by ensuring that zebu have access to mesic habitat. Additionally, the current interactions between pastoralism and riparian habitats should be studied to help conserve the island’s biodiversity.

Morphofunctional examination of the carpal bones of pygmy hippopotamus from Ayia Napa, Cyprus.

This study provides a complete morphological description and functional analysis of the carpal bones of the endemic pygmy hippopotamus Phanourios minor, derived from the Upper Pleistocene site of Ayia Napa. From this deposit, numerous skeletal remains of this fossil hippo have been collected, making the locality one of the richest in Cyprus. The carpal bones were compared with those of extant hippopotamuses, to determine the changes that occurred in the fossil hippo. Examination of the elements showed that Phanourios presented some important features that were common among the endemic fossil ungulates of the Mediterranean islands. The carpal bones display a proximal-distal compression due to shortening of the distal part of the leg, due to the new ecological island conditions. However, they are robust with rough areas for strong muscular and ligament insertions, providing stability to the carpal joints, and low speed movement to the animal. The great flexor capabilities, and the limitation in ulnar deviation of the carpus, indicate that P. minor had increased agility in the sagittal plane and restricted transverse movements, while the suggestion of a more unguligrade stance for the species is ambiguous. Thus, the endemic Cypriot hippos developed specialized locomotion, suitable for walking on the rugged terrain of Cyprus, which seems to be different from that of its extant counterparts.