Phylogenomics of Afrotherian mammals and improved resolution of extant Paenungulata.

Molecular investigations have gathered a diverse set of mammals-predominantly African natives like elephants, hyraxes, and aardvarks-into a clade known as Afrotheria. Nevertheless, the precise phylogenetic relationships among these species remain contentious. Here, we sourced orthologous markers and ultraconserved elements to discern the interordinal connections among Afrotherian mammals. Our phylogenetic analyses bolster the common origin of Afroinsectiphilia and Paenungulata, and propose Afrosoricida as the closer relative to Macroscelidea rather than Tubulidentata, while also challenging the notion of Sirenia and Hyracoidea as sister taxa. The approximately unbiased test and the gene concordance factor uniformly recognized the alliance of Proboscidea with Hyracoidea as the dominant topology within Paenungulata. Investigation into sites with extremly high phylogenetic signal unveiled their potential to intensify conflicts in the Paenungulata topology. Subsequent exploration suggested that incomplete lineage sorting was predominantly responsible for the observed contentious relationships, whereas introgression exerted a subsidiary influence. The divergence times estimated in our study hint at the Cretaceous-Paleogene (K-Pg) extinction event as a catalyst for Afrotherian diversification. Overall, our findings deliver a tentative but insightful overview of Afrotheria phylogeny and divergence, elucidating these relationships through the lens of phylogenomics.

Petrosal and bony labyrinth morphology of the stem paenungulate mammal (Paenungulatomorpha) Ocepeia daouiensis from the Paleocene of Morocco.

Based on high-resolution computed tomography, we describe in detail the petrosal and inner ear anatomy of one of the few known African stem paenungulates (Paenungulatomorpha), Ocepeia daouiensis from the Selandian of the Ouled Abdoun phosphate basin (Morocco). The petrosal of Ocepeia displays some remarkable, probably derived features (among eutherians) such as relatively small pars cochlearis, pars canalicularis labyrinth (including small semicircular canals), a large wing-like pars mastoidea, a large and inflated tegmen tympani, and the dorsoventral orientation of the large canal for the ramus superior. The presence of small semicircular canals in Ocepeia is an interesting shared trait with tenrecoidean afrotherians. Otherwise, and consistent with a general primitive skull morphology, the middle ear and labyrinth of Ocepeia daouiensis is characterised by many plesiomorphic traits close to the eutherian generalised plan. This adds to the rather generalised morphology of the earliest crown paenungulates such as Eritherium, Phosphatherium and Seggeurius to support an ancestral paenungulatomorph morphotype poorly derived from the eutherian pattern. As a result, Ocepeia provides key morphological and fossil data to test phylogenetic relationships of the Afrotheria (including Paenungulatomorpha) at the placental root mostly inferred from molecular studies.

Macronutrient composition of milk from two captive African elephant (Loxodonta africana) cows.

We assayed 31 milk samples collected from two African elephant cows housed at the Indianapolis Zoo across lactation (birth to calf age 973 days) for macronutrient composition (water, fat, protein, sugar, gross energy [GE], ash, calcium, and phosphorus). All assays were performed at the Smithsonian National Zoological Park Nutrition Laboratory, Washington, DC (SNZP) using standard methods developed at SNZP. Milk constituents are expressed on a weight-per-weight basis (%) and as a proportion each constituent contributes to milk energy. Calf weights were recorded, and growth rate calculated. The macronutrient composition of the African elephant milk samples was compared to previously published results for Asian elephants using analysis of covariance. African elephant milk is similar to Asian elephant milk, being moderately high in fat and energy and low in sugar. The mean values across lactation (excluding colostrum; n = 28) are 5.6 ± 0.3% crude protein, 3.1 ± 0.3% sugar, 13.0 ± 1.0% fat, and GE of 1.63 ± 0.10 kcal/g. Milk composition did not differ between cows. Milk composition significantly changed over lactation; fat and protein increased, and sugar decreased with calf age, comparable to previously reported data for African and Asian elephant milk. The proportion of milk energy from fat increased and that from sugar decreased over lactation, but the energy from protein was relatively constant. Protein contributed a higher proportion of energy to African elephant milk compared to Asian elephant milk (20.6% vs. 17.0%, p = .001). Despite this, calf growth rate was similar between the species, with the calves in this study gaining about 0.8 kg/day for the first 6 months.

Macronutrient composition of longitudinal milk samples from captive aardvarks (Orycteropus afer).

Hand-rearing and assisted-rearing aardvarks in captivity has become commonplace and has led to success in breeding the species. However, the macronutrient content of aardvark milk past 1 month of age is unknown. A better understanding of aardvark milk composition would enhance captive management efforts. Here, we assayed milk samples from two captive individuals from 2 to 114 days postpartum (N = 21) for dry matter, fat, crude protein, total sugar, ash (total minerals), calcium (Ca), phosphorus (P), and gross energy. The body weight of one calf was measured from birth to weaning. Milk macronutrient composition was compared to that of other Afrotherian species and Xenarthra species with similar diets. Average protein, fat, and sugar concentrations of aardvark milk across lactation were 12.3%, 13.6%, and 2.5%, respectively. Ash averaged 1.9%, with Ca (0.50%) and P (0.35%) accounting for about 45% of total minerals. All measured nutrients increased over lactation except sugar, which decreased. Aardvark milk is high in energy (2.12 kcal/g) mostly derived from fat and protein and little energy from sugar. Calf growth was linear (r2 = 0.995) with a mean gain of 159 g/day, achieving almost 30% of adult weight at weaning. Within Afrotheria, aardvark milk is higher in fat and protein and lower in sugar than elephant milk and more closely resembles the milk of its fellow insectivore, the elephant shrew. Aardvark milk is also similar in composition to milk of insectivorous Xenarthra species (nine-banded armadillo and giant anteater). Aardvark milk composition is consistent with the species' high-protein diet, fast growth, and nursing pattern.

Life on the rocks: Multilocus phylogeography of rock hyrax (Procavia capensis) from southern Africa.

Understanding the role of geography and climatic cycles in determining patterns of biodiversity is important in comparative and evolutionary biology and conservation. We studied the phylogeographic pattern and historical demography of a rock-dwelling small mammal species from southern Africa, the rock hyrax Procavia capensis capensis. Using a multilocus coalescent approach, we assessed the influence of strong habitat dependence and fluctuating regional climates on genetic diversity. We sequenced a mitochondrial gene (cytochrome b) and two nuclear introns (AP5, PRKC1) supplemented with microsatellite genotyping, in order to assess evolutionary processes over multiple temporal scales. In addition, distribution modelling was used to investigate the current and predicted distribution of the species under different climatic scenarios. Collectively, the data reveal a complex history of isolation followed by secondary contact shaping the current intraspecific diversity. The cyt b sequences confirmed the presence of two previously proposed geographically and genetically distinct lineages distributed across the southern African Great Escarpment and north-western mountain ranges. Molecular dating suggests Miocene divergence of the lineages, yet there are no discernible extrinsic barriers to gene flow. The nuclear markers reveal incomplete lineage sorting or ongoing mixing of the two lineages. Although the microsatellite data lend some support to the presence of two subpopulations, there is weak structuring within and between lineages. These data indicate the presence of gene flow from the northern into the southern parts of the southern African sub-region likely following the secondary contact. The distribution modelling predictably reveal the species' preference for rocky areas, with stable refugia through time in the northern mountain ranges, the Great Escarpment, as well as restricted areas of the Northern Cape Province and the Cape Fold Mountains of South Africa. Different microclimatic variables appear to determine the distributional range of the species. Despite strong habitat preference, the micro-habitat offered by rocky crevices and unique life history traits likely promoted the adaptability of P. capensis, resulting in the widespread distribution and persistence of the species over a long evolutionary period. Spatio-temporal comparison of the evolutionary histories of other co-distributed species across the rocky landscapes of southern Africa will improve our understanding of the regional patterns of biodiversity and local endemism.

Multiple Loci and Complete Taxonomic Sampling Resolve the Phylogeny and Biogeographic History of Tenrecs (Mammalia: Tenrecidae) and Reveal Higher Speciation Rates in Madagascar's Humid Forests.

The family Tenrecidae (tenrecs) is one of only four extant terrestrial mammal lineages to have colonized and diversified on Madagascar. Over the last 15 years, several studies have disagreed on relationships among major tenrec lineages, resulting in multiple reinterpretations of the number and timing of historical transoceanic dispersal events between Africa and Madagascar. We reconstructed the phylogeny of Tenrecidae using multiple loci from all recognized extant species and estimated divergence timing using six fossil calibrations within Afrotheria. All phylogenetic analyses strongly support monophyly of the Malagasy tenrecs, and our divergence timing analysis places their colonization of the island at 30-56 Ma. Our comprehensive phylogeny supports three important taxonomic revisions that reflect the evolutionary history of tenrecs: (1) we formally elevate the African otter shrews to their own family Potamogalidae, thereby rendering extant Tenrecidae entirely endemic to Madagascar; (2) we subsume the semiaquatic genus Limnogale within the shrew tenrec genus Microgale; and (3) we re-elevate the two largest-bodied shrew tenrecs, Microgale dobsoni and Microgale talazaci, to the genus Nesogale Thomas (1918) Finally, we use recently summarized habitat data to test the hypothesis that diversification rates differ between humid and arid habitats on Madagascar, and we compare three common methods for ancestral biogeographic reconstruction. These analyses suggest higher speciation rates in humid habitats and reveal a minimum of three and more likely five independent transitions to arid habitats. Our results resolve the relationships among previously recalcitrant taxa, illuminate the timing and mechanisms of major biogeographic patterns in an extraordinary example of an island radiation, and permit the first comprehensive, phylogenetically consistent taxonomy of Madagascar's tenrecs.

The ear region of earliest known elephant relatives: new light on the ancestral morphotype of proboscideans and afrotherians.

One of the last major clades of placental mammals recognized was the Afrotheria, which comprises all main endemic African mammals. This group includes the ungulate-like paenungulates, and among them the elephant order Proboscidea. Among afrotherians, the petrosal anatomy remains especially poorly known in Proboscidea. We provide here the first comparative CT scan study of the ear region of the two earliest known proboscideans (and paenungulates), Eritherium and Phosphatherium, from the mid Palaeocene and early Eocene of Morocco. It is helpful to characterize the ancestral morphotype of Proboscidea to understand petrosal evolution within proboscideans and afrotherians. The petrosal structure of these two taxa shows several differences. Eritherium is more primitive than Phosphatherium and closer to the basal paenungulate Ocepeia in several traits (inflated tegmen tympani, very deep fossa subarcuata and ossified canal for ramus superior of stapedial artery). Phosphatherium, however, retains plesiomorphies such as a true crus commune secundaria. A cladistic analysis of petrosal traits of Eritherium and Phosphatherium among Proboscidea results in a single tree with a low level of homoplasy in which Eritherium, Phosphatherium and Numidotherium are basal. This contrasts with previous phylogenetic studies showing homoplasy in petrosal evolution among Tethytheria. It suggests that evolutionary modalities of petrosal characters differ with the taxonomic level among Afrotheria: noticeable convergences occurred among the paenungulate orders, whereas little homoplasy seems to have occurred at intra-ordinal level in orders such as Proboscidea. Most petrosal features of both Eritherium and Phosphatherium are primitive. The ancestral petrosal morphotype of Proboscidea was not specialized but was close to the generalized condition of paenungulates, afrotherians, and even eutherians. This is consistent with cranial and dental characters of Eritherium, suggesting that the ancestral morphotypes of the different paenungulate orders were close to each other. Specializations occurred rapidly after the ordinal radiation of Paenungulata.

Fossils and living taxa agree on patterns of body mass evolution: a case study with Afrotheria.

Most of life is extinct, so incorporating some fossil evidence into analyses of macroevolution is typically seen as necessary to understand the diversification of life and patterns of morphological evolution. Here we test the effects of inclusion of fossils in a study of the body size evolution of afrotherian mammals, a clade that includes the elephants, sea cows and elephant shrews. We find that the inclusion of fossil tips has little impact on analyses of body mass evolution; from a small ancestral size (approx. 100 g), there is a shift in rate and an increase in mass leading to the larger-bodied Paenungulata and Tubulidentata, regardless of whether fossils are included or excluded from analyses. For Afrotheria, the inclusion of fossils and morphological character data affect phylogenetic topology, but these differences have little impact upon patterns of body mass evolution and these body mass evolutionary patterns are consistent with the fossil record. The largest differences between our analyses result from the evolutionary model, not the addition of fossils. For some clades, extant-only analyses may be reliable to reconstruct body mass evolution, but the addition of fossils and careful model selection is likely to increase confidence and accuracy of reconstructed macroevolutionary patterns.

Less is more in mammalian phylogenomics: AT-rich genes minimize tree conflicts and unravel the root of placental mammals.

Despite the rapid increase of size in phylogenomic data sets, a number of important nodes on animal phylogeny are still unresolved. Among these, the rooting of the placental mammal tree is still a controversial issue. One difficulty lies in the pervasive phylogenetic conflicts among genes, with each one telling its own story, which may be reliable or not. Here, we identified a simple criterion, that is, the GC content, which substantially helps in determining which gene trees best reflect the species tree. We assessed the ability of 13,111 coding sequence alignments to correctly reconstruct the placental phylogeny. We found that GC-rich genes induced a higher amount of conflict among gene trees and performed worse than AT-rich genes in retrieving well-supported, consensual nodes on the placental tree. We interpret this GC effect mainly as a consequence of genome-wide variations in recombination rate. Indeed, recombination is known to drive GC-content evolution through GC-biased gene conversion and might be problematic for phylogenetic reconstruction, for instance, in an incomplete lineage sorting context. When we focused on the AT-richest fraction of the data set, the resolution level of the placental phylogeny was greatly increased, and a strong support was obtained in favor of an Afrotheria rooting, that is, Afrotheria as the sister group of all other placentals. We show that in mammals most conflicts among gene trees, which have so far hampered the resolution of the placental tree, are concentrated in the GC-rich regions of the genome. We argue that the GC content-because it is a reliable indicator of the long-term recombination rate-is an informative criterion that could help in identifying the most reliable molecular markers for species tree inference.

Afrotheria genome; overestimation of genome size and distinct chromosome GC content revealed by flow karyotyping.

Afrotheria genome size is reported to be over 50% larger than that of human, but we show that this is a gross overestimate. Although genome sequencing in Afrotheria is not complete, extensive homology with human has been revealed by chromosome painting. We provide new data on chromosome size and GC content in four Afrotherian species using flow karyotyping. Genome sizes are 4.13 Gb in aardvark, 4.01 Gb in African elephant, 3.69 Gb in golden mole and 3.31 Gb in manatee, whereas published results show a mean of 5.18 Gb for Afrotheria. Genome GC content shows a negative correlation with size, indicating that this is due to differences in the amount of AT-rich sequences. Low genome GC content and small variance in chromosome GC content are characteristic of aardvark and elephant and may be associated with the high degree of conserved synteny, suggesting that these are features of the Afrotherian ancestral genome.

Cytochemical staining of leukocytes and platelets in the Florida manatee (Trichechus manatus latirostris): identification of a bilobed monocyte similar to other members of the Paenungulata.

Manatees (Antillean-, Amazonian, and African-) and dugongs belong to the Order Sirenia, and when combined with elephants and rock hyraxes, form the Paenungulata. A bilobed mononuclear cell has previously been identified in elephants and rock hyraxes, but not in manatees and dugongs, with cytochemical staining identifying these cells as bilobed monocytes in elephants. The objective of this study was to characterize leukocytes (white blood cells, WBC) and platelets in blood films of Florida manatees (Trichechus manatus latirostris; n = 8) using one routine hematological (Wright-Giemsa) and eight cytochemical stains: alkaline phosphatase (ALP), α-naphthyl butyrate esterase (ANBE), chloroacetate esterase (CAE), Luna, myeloperoxidase (MPx), periodic acid-Schiff (PAS), Sudan black B (SBB), and toluidine blue (TB). Heterophils and lymphocytes comprised most of the WBC, with low numbers of eosinophils, basophils, and monocytes. Additionally, 1-3% of the WBC were bilobed mononuclear cells. Bilobed mononuclear cell proportions were similar to rock hyraxes, but lower than elephants (approximate range 20-60%). Heterophils and eosinophils were positive for MPx, ALP, SBB, and PAS, with heterophils also being positive for CAE. Most of the lymphocytes were positive for ANBE and they were variably positive for CAE. Monocytes and bilobed mononuclear cells had similar cytochemical staining reactions (variably positive for all stains, except Luna and TB), supporting a monocytic origin, like elephants. Platelets were ANBE- and PAS-positive. Luna stain was useful for identifying eosinophils and TB was uninformative. This study provides new information on the morphological features and cytochemical staining characteristics of WBC and platelets and will aid in obtaining accurate hematological data of Florida manatees.

Cholinergic, catecholaminergic, serotonergic, and orexinergic neuronal populations in the brain of the lesser hedgehog tenrec (Echinops telfairi).

The current study provides an analysis of the cholinergic, catecholaminergic, serotonergic, and orexinergic neuronal populations, or nuclei, in the brain of the lesser hedgehog tenrec, as revealed with immunohistochemical techniques. For all four of these neuromodulatory systems, the nuclear organization was very similar to that observed in other Afrotherian species and is broadly similar to that observed in other mammals. The cholinergic system shows the most variation, with the lesser hedgehog tenrec exhibiting palely immunopositive cholinergic neurons in the ventral portion of the lateral septal nucleus, and the possible absence of cholinergic neurons in the parabigeminal nucleus and the medullary tegmental field. The nuclear complement of the catecholaminergic, serotonergic and orexinergic systems showed no specific variances in the lesser hedgehog tenrec when compared to other Afrotherians, or broadly with other mammals. A striking feature of the lesser hedgehog tenrec brain is a significant mesencephalic flexure that is observed in most members of the Tenrecoidea, as well as the closely related Chrysochlorinae (golden moles), but is not present in the greater otter shrew, a species of the Potomogalidae lineage currently incorporated into the Tenrecoidea. In addition, the cholinergic neurons of the ventral portion of the lateral septal nucleus are observed in the golden moles, but not in the greater otter shrew. This indicates that either complex parallel evolution of these features occurred in the Tenrecoidea and Chrysochlorinae lineages, or that the placement of the Potomogalidae within the Tenrecoidea needs to be re-examined.

Uncoupling elephant TP53 and cancer.

Elephant testicles do not descend, with implications for sperm production being hot enough to compromise germline DNA replication/repair. Uniquely, elephants also possess 20 copies of a gene encoding for the p53 protein. Did elephants evolve multiplication of the TP53 gene complex to protect their germline rather than to fight cancer?

A Comparative Genomic and Phylogenetic Investigation of the Xenobiotic Metabolism Enzymes of Cytochrome P450 in Elephants Shows Loss in CYP2E and CYP4A.

Cytochrome P450 is an important enzyme that metabolizes a variety of chemicals, including exogenous substances, such as drugs and environmental chemicals, and endogenous substances, such as steroids, fatty acids, and cholesterol. Some CYPs show interspecific differences in terms of genetic variation. As little is known about the mechanisms of elephant metabolism, we carried out a comparative genomic and phylogenetic analysis of CYP in elephants. Our results suggest that elephant CYP genes have undergone independent duplication, particularly in the CYP2A, CYP2C, and CYP3A genes, a unique cluster specific to elephant species. However, while CYP2E and CYP4A were conserved in other Afrotheria taxa, their decay in elephants resulted in genetic dysfunction (pseudogene). These findings outline several remarkable characteristics of elephant CYP1-4 genes and provide new insights into elephant xenobiotic metabolism. Further functional investigations are necessary to characterize elephant CYP, including expression patterns and interactions with drugs and sensitivities to other chemicals.

Review of sensory modalities of sirenians and the other extant Paenungulata clade.

Extant members of Paenungulata (sirenians, proboscideans, and hyracoideans) form a monophyletic clade which originated in Africa. While paenungulates are all herbivorous, they differ greatly in size, life history, and habitat. Therefore, we would expect both phylogenetically related similarities and ecologically driven differences in their use and specializations of sensory systems, especially in adaptations in sirenians related to their fully aquatic habitat. Here we review what is known about the sensory modalities of this clade in an attempt to better elucidate their sensory adaptations. Manatees have a higher frequency range for hearing than elephants, who have the best low-frequency hearing range known to mammals, while the hearing range of hyraxes is unknown. All paenungulates have vibrissae assisting in tactile abilities such as feeding and navigating the environment and share relatively small eyes and dichromatic vision. Taste buds are present in varying quantities in all three orders. While the olfactory abilities of manatees and hyraxes are unknown, elephants have an excellent sense of smell which is reflected by having the relatively largest cranial nerve related to olfaction among the three lineages. Manatees have the relatively largest trigeminal nerve-the nerve responsible for, among other things, mystacial vibrissae-while hyraxes have the relatively largest optic nerve (and therefore, presumably, the best vision) among the Paenungulata. All three orders have diverged significantly; however, they still retain some anatomical and physiological adaptations in common with regard to sensory abilities.

Total evidence time-scaled phylogenetic and biogeographic models for the evolution of sea cows (Sirenia, Afrotheria).

Molecular phylogenetic studies that have included sirenians from the genera Trichechus, Dugong, and Hydrodamalis have resolved their interrelationships but have yielded divergence age estimates that are problematically discordant. The ages of these lineage splits have profound implications for how to interpret the sirenian fossil record-including clade membership, biogeographic patterns, and correlations with Earth history events. In an effort to address these issues, here we present a total evidence phylogenetic analysis of Sirenia that includes living and fossil species and applies Bayesian tip-dating methods to estimate their interrelationships and divergence times. In addition to extant sirenians, our dataset includes 56 fossil species from 106 dated localities and numerous afrotherian outgroup taxa. Genetic, morphological, temporal, and biogeographic data are assessed simultaneously to bring all available evidence to bear on sirenian phylogeny. The resulting time-tree is then used for Bayesian geocoordinates reconstruction analysis, which models ancestral geographic areas at splits throughout the phylogeny, thereby allowing us to infer the direction and timing of dispersals. Our results suggest that Pan-Sirenia arose in North Africa during the latest Paleocene and that the Eocene evolution of stem sirenians was primarily situated in the Tethyan realm. In the late Eocene, some lineages moved into more northern European latitudes, an area that became the source region for a key trans-Atlantic dispersal towards the Caribbean and northern-adjacent west Atlantic. This event led to the phylogenetic and biogeographic founding of crown Sirenia with the Dugongidae-Trichechidae split occurring at the Eocene-Oligocene boundary (~33.9 Ma), temporally coincident with the onset of dropping global sea levels and temperatures. This region became the nexus of sirenian diversification and supported taxonomically-rich dugongid communities until the earliest Pliocene. The Dugonginae-Hydrodamalinae split occurred near Florida during the early Miocene (~21.2 Ma) and was followed by a west-bound dispersal that gave rise to the Pacific hydrodamalines. The late middle Miocene (~12.2 Ma) split of Dugong from all other dugongines also occurred near Florida and our analyses suggest that the Indo-Pacific distribution of modern dugongs is the result of a trans-Pacific dispersal. From at least the early Miocene, trichechid evolution was based entirely in South America, presumably within the Pebas Wetlands System. We infer that the eventual establishment of Amazon drainage into the South Atlantic allowed the dispersal of Trichechus out of South America no earlier than the mid-Pliocene. Our analyses provide a new temporal and biogeographic framework for understanding major events in sirenian evolution and their possible relationships to oceanographic and climatic changes. These hypotheses can be further tested with the recovery and integration of new fossil evidence.

Principles of 3D chromosome folding and evolutionary genome reshuffling in mammals.

Studying the similarities and differences in genomic interactions between species provides fertile grounds for determining the evolutionary dynamics underpinning genome function and speciation. Here, we describe the principles of 3D genome folding in vertebrates and show how lineage-specific patterns of genome reshuffling can result in different chromatin configurations. We (1) identified different patterns of chromosome folding in across vertebrate species (centromere clustering versus chromosomal territories); (2) reconstructed ancestral marsupial and afrotherian genomes analyzing whole-genome sequences of species representative of the major therian phylogroups; (3) detected lineage-specific chromosome rearrangements; and (4) identified the dynamics of the structural properties of genome reshuffling through therian evolution. We present evidence of chromatin configurational changes that result from ancestral inversions and fusions/fissions. We catalog the close interplay between chromatin higher-order organization and therian genome evolution and introduce an interpretative hypothesis that explains how chromatin folding influences evolutionary patterns of genome reshuffling.

Pervasive duplication of tumor suppressors in Afrotherians during the evolution of large bodies and reduced cancer risk.

The risk of developing cancer is correlated with body size and lifespan within species. Between species, however, there is no correlation between cancer and either body size or lifespan, indicating that large, long-lived species have evolved enhanced cancer protection mechanisms. Elephants and their relatives (Proboscideans) are a particularly interesting lineage for the exploration of mechanisms underlying the evolution of augmented cancer resistance because they evolved large bodies recently within a clade of smaller-bodied species (Afrotherians). Here, we explore the contribution of gene duplication to body size and cancer risk in Afrotherians. Unexpectedly, we found that tumor suppressor duplication was pervasive in Afrotherian genomes, rather than restricted to Proboscideans. Proboscideans, however, have duplicates in unique pathways that may underlie some aspects of their remarkable anti-cancer cell biology. These data suggest that duplication of tumor suppressor genes facilitated the evolution of increased body size by compensating for decreasing intrinsic cancer risk.

From the gigantic blue whale to the minuscule bumblebee bat, animals come in all shapes and sizes. Any species can develop cancer, but some are more at risk than others. In theory, if every cell has the same probability of becoming cancerous, then bigger animals should get cancer more often since they have more cells than smaller ones. Amongst the same species, this relationship is true: taller people and bigger dogs have a greater cancer risk than their smaller counterparts. Yet this correlation does not hold when comparing between species: remarkably large creatures, like elephants and whales, are not more likely to have cancer than any other animal. But how have these gigantic animals evolved to be at lower risk for the disease? To investigate, Vazquez and Lynch compared the cancer risk and the genetic information of a diverse group of closely related animals with different body sizes. This included elephants, woolly mammoths and mastodons as well as their small relatives, the manatees, armadillos, and marmot-sized hyraxes. Examining these species' genomes revealed that, during evolution, elephants had acquired extra copies of 'tumour suppressor genes' which can sense and repair the genetic and cellular damages that turn healthy cells into tumours. This allowed the species to evolve large bodies while lowering their risk of cancer. Further studies could investigate whether other gigantic animals evolved similar ways to shield themselves from cancer; these could also examine precisely how having additional copies of cancer-protecting genes helps reduce cancer risk, potentially paving the way for new approaches to treat or prevent the disease.

Microscopic structure of the tongue in the lesser hedgehog tenrec (Echinops telfairi, Afrosoricida) and its relation to phylogenesis.

The tongue of the lesser hedgehog tenrec (Echinops telfairi) was evaluated by light and scanning electron microscopy. Dorsal and lateral surfaces of the tongue were densely covered with various types of lingual papillae. Two types of mechanical papillae (filiform and conical) and two types of sensory papillae (fungiform and vallate) were observed. Rostrocaudally, the number of spikes in filiform papillae increased. Filiform, crown-like papillae were found on the radix, as well as large conical papillae with a wide base and a caudally bent single spike. Fungiform papillae were scattered in four rows on the dorsal surface. Three vallate papillae in inverse triangular layout were developed in the area of the lingual radix. Taste buds were found in the sensory papillae, i.e., fungiform and vallate. The lingual glands were predominantly seromucous. These glands were evenly scattered from the caudal portion of the lingual apex to the lingual radix. Purely serous glands were localized close to vallate papillae. A histochemical analysis of glands was performed. Results were compared to the Afrotheria (tenrec's relatives) and Eulipotyphla (animals with similar feeding habits to tenrecs). This study is the first description of the lingual morphology in the member of the Order Afrosoricida. Moreover, it compares the effect of the phylogenetic relation to the effect of the feeding habits on the lingual morphology.

The hypercholinergic brain of the Cape golden mole (Chrysochloris asiatica).

Studies detailing the anatomy of the brain of the golden moles are few. A recent study indicated that in the Hottentot golden mole (a member of the Amblysominae clade), there was a broad, atypical, distribution of cholinergic interneurons in the olfactory bulb, cerebral cortex, hippocampus and amygdala. To determine whether this broad distribution of cholinergic neurons is shared by other species of golden mole, we here examine the brain of the Cape golden mole (a member of the Chrysochlorinae clade, representing the second major clade within the family Chrysochloridae). Our analyses indicates the presence of a similar widespread distribution of cholinergic interneurons in the Cape golden mole. Thus, we conclude that these features are derived morphological traits in the brains of golden moles. In addition, we describe the nuclei generally considered to be part of the typical cholinergic system in mammals. Whereas the vast majority of these generally reported cholinergic nuclei were the same as recorded in other Eutherian mammals, it was noted that the cholinergic nuclei involved in oculomotion were substantially reduced in size, or absent in the case of the abducens nucleus. In addition, there was an absence of the cholinergic medial septal nucleus, but the presence of a cholinergic lateral septal nucleus. The laterodorsal and pedunculopontine tegmental nuclei evince regions where the cholinergic neurons are densely packed. These are atypical features of the mammalian cholinergic system, which when combined with the widespread atypical distribution of cholinergic interneurons, reveals a family-specific complement of cholinergic nuclei in the Chrysochloridae.