Comparative study of the ciliary body and iris morphology in the anterior eye chamber of five different vertebrate classes.

One crucial component of the optical system is the ciliary body (CB). This body secretes the aqueous humour, which is essential to maintain the internal eye pressure as well as the clearness of the lens and cornea. The histological study was designed to provide the morphological differences of CB and iris in the anterior eye chambers of the following vertebrate classes: fish (grass carp), amphibians (Arabian toad), reptiles (semiaquatic turtle, fan-footed gecko, ocellated skink, Egyptian spiny-tailed lizard, Arabian horned viper), birds (common pigeon, common quail, common kestrel), and mammals (BALB/c mouse, rabbit, golden hamster, desert hedgehog, lesser Egyptian jerboa, Egyptian fruit bat). The results showed distinct morphological appearances of the CB and iris in each species, ranging from fish to mammals. The present comparative study concluded that the morphological structure of the CB and iris is the adaptation of species to either their lifestyle or survival in specific habitats.

Juvenile European hedgehogs (Erinaceus europaeus) at rescue centers and their release rate depending on their weight on admission.

This study aimed to assess the numbers of juvenile European hedgehogs admitted to rescue centers in the Czech Republic from the viewpoint of their weight on admission, the reason for their admission, and the success rate of their release back into the wild. The results of our study show varying levels of success in the rearing of hedgehogs admitted at different ages (weights) and a varying period required for their rehabilitation. The greatest chance of release was seen in hedgehogs with a weight on the admission of 500-599 g (64.22% released) and 400-499 g (63.31% released). In contrast, the smallest number of young hedgehogs successfully rehabilitated and released was seen in hoglets weighing 200-299 g (35.24% released) on admission, which corresponds to the weight of hedgehogs at the time of weaning. Time spent at a rescue center may pose an undesirable threat to the lives of animals in some categories. Hedgehogs weighing up to 99 g on admission spent the longest period time at rescue centers (a median of 48 days), while hedgehogs weighing 500-599 g on admission spent the shortest time (a median of 7 days). The majority of hedgehogs in the lowest weight categories were admitted due to their inability to survive on their own. A large percentage of hedgehogs of greater weight, in contrast, were juvenile hedgehogs brought to rescue centers needlessly. The percentage of released animals did not exceed 65%, however, even for entirely independent categories of older juveniles. From this perspective, the fact that hedgehogs are often brought to rescue centers in the belief that they are not self-sufficient young, though they are actually juvenile or even adult individuals that do not require human care, can be considered a significant finding.

Red Fluorescence of European Hedgehog (Erinaceus europaeus) Spines Results from Free-Base Porphyrins of Potential Microbial Origin.

Bioluminescence has been recognized as an important means for inter- and intra-species communication. A growing number of reports of red fluorescence occurring in keratinaceous materials have become available. The fluorophore(s) in these cases were shown to be, or suspected to be, free base porphyrins. The red fluorescence found in the downs of bustards was associated with inter-species signaling in mate selection. First reported in 1925, we confirm that spines of the European hedgehog (Erinaceus europaeus) when irradiated with UV (365-395 nm) light display red fluorescence localized in the light-colored sections of their proximal ends. Using reflectance fluorescence spectroscopy, we confirmed that the fluorophores responsible for the emission are free-base porphyrins, as suspected in the original report. Base-induced degradation of the spine matrix and subsequent HPLC, UV-vis, and ESI+ mass spectrometry analysis revealed the presence of a mixture of coproporphyrin III and uroporphyrin III as predominant porphyrins and a minor fraction of protoporphyrin IX. Investigation of the spine microbiome uncovered the abundant presence of bacteria known to secrete and/or interconvert porphyrins and that are not present on the non-fluorescing quills of the North American porcupine (Erethizon dorsatum). Given this circumstantial evidence, we propose the porphyrins could originate from commensal bacteria. Furthermore, we hypothesize that the fluorescence may be incidental and of no biological function for the hedgehog.

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.

Histological characterisation of the skin of the Paraechinus hypomelas, Brandt, 1836 (Erinaceidae: Eulipotyphla).

BACKGROUND: The current study represents the first description of the histological characterisations of the normal skin of Brandt's hedgehog (paraechinus hypomelas). MATERIALS AND METHODS: Skin samples were collected from abdomen, back, nostril and cloacal regions. RESULTS: The skin consisted of 3 layers including epidermis, dermis and hypodermis. The epidermis was covered by a layer of keratinised squamous epithelium mainly in the back region, but the skin keratinisation was present with a little amount or may was absent in other regions. Histologically, the total thickness of skin was maximum on the back and minimum on the cloacal regions. The epidermis consisted of 4 layers and stratum lucidum was absent in all regions. Beneath the epidermis, the dermis layer was constituted of dense connective tissue in which the hair follicles, sweat glands, sebaceous glands, arrector pilli muscles and blood vessels were present. The sweat and sebaceous glands were more populated in the nostril region. The hair follicles were located in the epidermal and dermal regions. Vibrissae were only in the nostrils region and characterised from other hairs by their large and well innervated hair follicle which was surrounded by the blood sinus. CONCLUSIONS: The present findings show that in Brandt's hedgehog (paraechinus hypomelas) the back and cloacal regions have thickest and thinnest skin respectively as compared to the nostril and abdominal regions. In addition, sebaceous and sweat glands were mainly populated in the nostril region.

Morphology of the Lingual Papillae of the Japanese lesser flying squirrel and four-toed hedgehog.

Author examined the dorsal lingual surfaces of the adult Japanese lesser flying squirrel (Pteromys momonga) and four-toed hedgehog (Atelerix albiventris) by scanning electron microscopy. In the Japanese lesser flying squirrel, the filiform papilla of the lingual body consisted of a large conical papilla. The filiform papilla of the lingual prominence was spoon in shape. The fungiform papillae were round in shape and scattered among the filiform papillae. Many foliate papillae were observed on the posterolateral regions of the lingual body. The foliate papillae had some ridges separated by deep grooves. The vallate papilla was located between lingual body and root. Several long conical papillae derived from the posterolateral margin of the tongue. In the four-toed hedgehog, the filiform papilla of the lingual apex had a conical process. The filiform papilla of the lingual body had some processes. The fungiform papillae were round in shape. The foliate papillae were observed on the posterolateral regions of the lingual body. The papilla was separated from each other by a furrow. The vallate papilla consisted of a central papilla and an annular pad. These findings suggest that in the structure of the lingual papillae of the Japanese lesser flying squirrel there is similar to that of the sugar glider and the lingual papillae of the four-toed hedgehog is different from that of the Japanese lesser flying squirrel.

Descriptive studies on the tongue of two micro-mammals inhabiting the Egyptian fauna; the Nile grass rat (Arvicanthis niloticus) and the Egyptian long-eared hedgehog (Hemiechinus auritus).

The current study aimed to describe the anatomical features of the tongues of two micro-mammals common in the Egyptian fauna; the Nile grass rat (Arvicathis niloticus), and the Egyptian long-eared hedgehog (Hemiechinus auritus). The tongues of five adult individuals of each species were excised and processed histologically, histochemically, and morphometrically. Statistical analysis comparing the relative tongue length in both species showed that there was a significant difference, which may correlate with the difference in feeding preferences. Grossly, the Nile grass rat has a dorsal lingual prominence with bifurcated apex while, the long-eared hedgehog has a median slight elevation with rounded apex. Numerous forms of mechanical and gustatory papillae are scattered along the lingual dorsal epithelium. The histochemical detection of keratin by Holland's trichrome stain showed an intense expression in the case of A. niloticus and mild expression in H. auritus. The framework of the tongue (entoglossum) is supported by either a core of cartilage in H. auritus or bone in A. niloticus which incorporated in the lingual root. The lingual glands also showed marked variation, the Nile grass rat exhibit dense populations of mucous-secreting glands and lesser populations of serous-secreting glands, the contrary is true in H. auritus. In conclusion, the micro and macro-anatomical features of the tongues of both species showed adaptive changes to accommodate the feeding lifestyle. Such type of studies using mammals from different phylogenetic traits and almost have different feeding preferences provide answers to many research questions related to tongue evolution among mammalian vertebrates.

Taxonomic revision of the genus Mesechinus (Mammalia: Erinaceidae) with description of a new species.

Hedgehogs in the genus Mesechinus (Family Erinaceidae), which include two currently recognized species (M. dauuricus and M. hughi), are distributed from northeast Mongolia to the upper Amur Basin in Russia and adjacent areas in northeast and northern China. In recent years, a population of Mesechinus hedgehogs was discovered from Mt. Gaoligong, southwestern Yunnan, China, far from the known distribution range of the genus. Furthermore, these hedgehogs are the only known population to be distributed at elevations higher than 2 100 m and in sympatry with gymnures. To evaluate the taxonomic status of these hedgehogs, we examined specimens representing Mesechinus taxa in China and further conducted morphometric and karyotypic analyses. Our results supported the existence of four species in China. Specifically, we identified the hedgehogs from Mt. Gaoligong as a new species, Mesechinus wangi sp. nov., and recognized M. miodon, previously considered as a synonym of either M. dauuricus or M. hughi, as a distinct species. Interestingly, we observed a supernumerary M4 on all specimens of Mesechinus wangi sp. nov., which is an extremely rare event in the evolution of mammalian dentition.

The gross anatomy of the male reproductive system of the European hedgehog (Erinaceus Europaeus).

Hedgehogs are small spiny-coated insectivores. Due to their low body weight, calm character, and easy maintenance, they are kept as pets. It is therefore worthwhile to care about hedgehogs' health problems and to provide pet owners with information about their reproduction. Moreover, it is necessary to be familiar with their anatomy so as to satisfy the need to improve nutrition and medical care, even surgery. This study was carried out on five adult male European hedgehogs euthanased in a chloroform chamber. The European hedgehog's oval testes are invisible in inguinal region because they have no true scrotal sac. The testes are located in the craniocaudal direction with dorsolateral epididymal attachments. The vesicular glands, the European hedgehog's largest accessory sex glands, are lobulated structures containing dorsomedial and ventrolateral parts on each side. The prostate is an oval gland with right and left lobes. The paired bulbourethral glands are laid on the ischiocavernosus muscle. Histologically the vesicular, prostate gland ducts and ductus deferens as well as urethra separately were discharged in a diverticlum at the level of the pelvic urethra end. A sigmoid flexure exists in the proximal part of shaft body of the penis. There are two retractor penile muscles. In dorsal end of the penile glans, there is a small urethral process with two nail- -like, needle-shaped structures. They are on both sides of the urethral process. Furthermore, there are two intromittent sacs (Sacculus urethralis) in the ventral part of the end of the penis. (Folia Morphol 2018; 77, 1: 36-43).

Morphological study of the European hedgehog (Erinaceus europaeus) tongue by SEM and LM.

The hedgehog tongue is a tactile and taste organ which carries out various functions. Detailed functional and morphological studies are required to clearly define the relationship of the hedgehog tongue with taste, food palatability, mastication and swallowing of food, as well as the production of sounds. The aim of this study was to determine the relationship between the morphological characteristics of the European hedgehog tongue and the lifestyle of this animal, as well as to compare findings with the results of studies on other vertebrates. Gross and micro-anatomical light and scanning electron microscopy studies revealed that the hedgehog tongue could be divided in three areas, namely the apex, body and root. A keratinized stratified squamous epithelium, which was smooth on the ventral surface but bore four types of papillae on the dorsal surface, lined the tongue. Three types of these papillae were found to have gustatory functions and to express their activity in close relation with the salivary glands. These simple conical filiform papillae were situated caudally and distributed one after the other without a break. The dome-shaped fungiform papillae on the apex, with the highest distribution rate on the apex edge, were small, but those on the body and root were large. The three circular vallate papillae were arranged in a triangular shape. The foliate papillae with a few tiny projections, found in a shallow furrow, were situated between the root and the body. Most of the nerve fibers observed in different sections of the tongue tissue were of the unmyelinated type, confirming that the main task of the hedgehog tongue was its gustatory function.

Anatomy and Disorders of the Oral Cavity of Miscellaneous Exotic Companion Mammals.

Unusual mammalian species such as the hedgehog, sugar glider, and miniature pig are encountered with increasing frequency in exotic companion medicine. Disease of the oral cavity can occur in any species; although occasionally encountered in exotic mammalian species, it is rarely described in the literature. Anatomy and dentition vary significantly; diagnosis and treatment are often extrapolated from that known in other species. The best-documented disease of the oral cavity in this group of species is oral neoplasia in the hedgehog.

The comparative gastrointestinal morphology of Jaculus jaculus (Rodentia) and Paraechinus aethiopicus (Erinaceomorpha).

Jaculus jaculus (Lesser Egyptian jerboa) and Paraechinus aethiopicus (Desert hedgehog) are small mammals which thrive in desert conditions and are found, among others, in the Arabian Peninsula. Jaculus jaculus is omnivorous while P. aethiopicus is described as being insectivorous. The study aims to describe the gastrointestinal tract (GIT) morphology of these animals which differ in diet and phylogeny. The GITs of J. jaculus (n = 8) and P. aethiopicus (n = 7) were weighed, photographed, and the length, basal surface areas, and luminal surface areas of each of the anatomically distinct gastrointestinal segments were determined. The internal aspects of each area were examined and photographed while representative histological sections of each area were processed to wax and stained using haematoxylin and eosin. Both species had a simple unilocular stomach which was confirmed as wholly glandular on histology sections. Paraechinus aethiopicus had a relatively simple GIT which lacked a caecum. The caecum of J. jaculus was elongated, terminating in a narrow cecal appendix which contained lymphoid tissue on histological examination. The internal aspect of the proximal colon of J. jaculus revealed distinct V-shaped folds. Stomach content analysis of J. jaculus revealed mostly plant and seed material and some insects, whereas P. aethiopicus samples showed plant material in addition to insects, indicating omnivorous feeding tendencies in areas where insects may be scarce.

Nuclear organisation of some immunohistochemically identifiable neural systems in five species of insectivore-Crocidura cyanea, Crocidura olivieri, Sylvisorex ollula, Paraechinus aethiopicus and Atelerix frontalis.

The organization of the cholinergic, catecholaminergic, and serotonergic neurons in the brains of five species of insectivores and the orexinergic (hypocretinergic) system in four insectivore species is presented. We aimed to investigate the nuclear complement of these neural systems in comparison to those of other mammalian species. Brains of insectivores were coronally sectioned and immunohistochemically stained with antibodies against choline acetyltransferase, tyrosine hydroxylase, serotonin and orexin-A. The majority of nuclei were similar among the species investigated and to mammals in general, but certain differences in the nuclear complement highlighted potential phylogenetic interrelationships. In the cholinergic system, the three shrew species lacked parabigeminal and Edinger-Westphal nuclei. In addition, the appearance of the laterodorsal tegmental nucleus in all insectivores revealed a mediodorsal arch. All three of these features are the same as those present in microchiropterans. The catecholaminergic system of the three shrew species lacked the A4 and A15d nuclei, as well as having an incipient A9v nucleus, again features found in microchiropteran brains. The serotonergic and orexinergic systems of the insectivores are similar to those seen across most eutherian mammals. The analysis of similarities and differences across mammalian species indicates a potential phylogenetic relationship between the Soricidae (shrews) and the microchiropterans.

Multicentric epitheliotropic T-cell lymphoma in an African hedgehog (Atelerix albiventris).

A 2-year-old female African hedgehog was presented with a 5-month history of pruritus, and diffuse spine and hair loss. A dermatologic examination revealed erythema, excoriation, scales, and crusting affecting the face, flanks, forelimbs, hindlimbs, and dorsal and ventral abdomen. Fine-needle aspiration was performed and skin biopsies were taken from several lesions for cytologic and histologic evaluation. The aspirates yielded smears characterized by a monomorphic population of medium-sized to large lymphocytes with scant to moderate amounts of clear to moderately basophilic cytoplasm and distinct nucleoli along with a low number of cytoplasmic fragments. On histopathologic examination, there were dense dermal lymphoid infiltrates invading the dermis and a monomorphic population of round cells that had infiltrated the overlying epidermis. Epitheliotropic cutaneous lymphoma was diagnosed based on morphologic features. Additional immunochemical analysis using anti-CD3 and anti-CD79a antibodies revealed strong CD3 expression by the tumor cells, which confirmed epitheliotropic cutaneous T-cell lymphoma. This is the first description of a multicentric pattern of epitheliotropic cutaneous T-cell lymphoma in an African hedgehog.

The placentation of eulipotyphla-reconstructing a morphotype of the Mammalian placenta.

Placentation determines the developmental status of the neonate, which can be considered as the most vulnerable stage in the mammalian life cycle. In this respect, the different evolutionary and ecological adaptations of marsupial and placental mammals have most likely been associated with the different reproductive strategies of the two therian clades. The morphotypes of marsupial and placental neonates, as well as the placental stem species pattern of Marsupialia, have already been reconstructed. To contribute to a better understanding of the evolution of Placentalia, a histological and ultrastructural investigation of the placenta in three representatives of Eulipotyphla, that is, core insectivores, has been carried out in this study. We studied the Musk shrew (Suncus murinus), the four-toed hedgehog (Atelerix albiventris), and the Iberian mole (Talpa occidentalis). As a result, a eulipotyphlan placental morphotype consisting of a compact and invasive placenta was reconstructed. This supports the widely accepted hypothesis that the stem lineage of Placentalia is characterized by an invasive, either endothelio- or hemochorial placenta. Evolutionary transformations toward a diffuse, noninvasive placenta occurred in the stem lineages of lower primates and cetartiodactyles and were associated with prolonged gestation and the production of few and highly precocial neonates. Compared to the choriovitelline placenta of Marsupialia, the chorioallantoic placenta of Placentalia allows for a more intimate contact and is associated with more advanced neonates.

Adult neurogenesis in the hedgehog (Erinaceus concolor) and mole (Talpa europaea).

We investigated adult neurogenesis in two species of mammals belonging to the superorder Laurasiatheria, the southern white-breasted hedgehog (order Erinaceomorpha, species Erinaceus concolor) from Armenia and the European mole (order Soricomorpha, species Talpa europaea) from Poland. Neurogenesis in the brain of these species was examined immunohistochemically, using the endogenous markers doublecortin (DCX) and Ki-67, which are highly conserved among species. We found that in both the hedgehog and mole, like in the majority of earlier investigated mammals, neurogenesis continues in the subventricular zone (SVZ) of the lateral ventricles and in the dentate gyrus (DG). In the DG of both species, DCX-expressing cells and Ki-67-labeled cells were present in the subgranular and granular layers. In the mole, a strong bundle of DCX-labeled processes, presumably axons of granule cells, was observed in the center of the hilus. Proliferating cells (expressing Ki-67) were identified in the SVZ of lateral ventricles of both species, but neuronal precursor cells (expressing DCX) were also observed in the olfactory bulb (OB). In both species, the vast majority of cells expressing DCX in the OB were granule cells with radially orientated dendrites, although some periglomerular cells surrounding the glomeruli were also labeled. In addition, this paper is the first to show DCX-labeled fibers in the anterior commissure of the hedgehog and mole. These fibers must be axons of new neurons making interhemispheric connections between the two OB or piriform (olfactory) cortices. DCX-expressing neurons were observed in the striatum and piriform cortex of both hedgehog and mole. We postulate that in both species a fraction of cells newly generated in the SVZ migrates along the rostral migratory stream to the piriform cortex. This pattern of migration resembles that of the 'second-wave neurons' generated during embryonal development of the neocortex rather than the pattern observed during development of the allocortex. In spite of the presence of glial cells alongside DCX-expressing cells, we never found colocalization of DCX protein with a glial marker (vimentin or glial fibrillary acidic protein).

Radiological investigations of the hedgehog (Erinaceus concolor) appendicular skeleton.

The normal radiographic anatomy of the healthy hedgehog can help to identify anatomic features unique to the hedgehog while comparing it with other small mammals, such as the dog and cat. Radiographic examination is a method that can play an important role in the diagnosis of a wide variety of skeletal diseases. Seven (2 males, 5 females) free-living hedgehogs (Erinaceus concolor) from the Urmia region of Iran were selected for this study. Lateral and craniocaudal radiographs from the front and hind limbs were obtained. The radiographs from these hedgehogs were compared with the normal canine and feline skeletal radiographic anatomy. On the forelimb radiographs, the clavicle was observed as a complete bone connected to the scapula and manubrium. There are three and five carpal bones in the proximal and distal rows, respectively, as in the dog and cat. The pelvis has a larger obturator foramen when compared with the dog and cat. In the lateral view, the pubis and ischium are relatively larger than in the dog and cat and have a more ventral position. The tarsal bones are similar to those of the dog and cat. The number of phalanges and sesamoid bones in the forelimb and hindlimb are likewise similar to those found in the dog and cat.

Pyramidal neurons in the septal and temporal CA1 field of the human and hedgehog tenrec hippocampus.

The present study examines comparatively the cellular density of disector-counted/Nissl-stained CA1 pyramidal neurons and the morphometric characteristics (dendritic number/length, spine number/density and Sholl-counted dendritic branch points/20 microm) of the basal and apical dendritic systems of Golgi-impregnated CA1 neurons, in the septal and temporal hippocampus of the human and hedgehog tenrec brain. The obtained results indicate that in both hippocampal parts the cellular density of the CA1 pyramidal neurons is lower in human than in tenrec. However, while the human pyramidal cell density is higher in the septal hippocampal part than in the temporal one, in the tenrec the density of these cells is higher in the temporal part. The dendritic tree of the CA1 pyramidal cells, more developed in the septal than in temporal hippocampus in both species studied, is in general more complex in the human hippocampus. The basal and the apical dendritic systems exhibit species related morphometric differences, while dendrites of different orders exhibit differences in their number and length, and in their spine density. Finally, in both species, as well as hippocampal parts and dendritic systems, changes of dendritic morphometric features along ascending dendritic orders fluctuate in a similar way, as do the number of dendritic branch points in relation to the distance from the neuron soma.

Cranial features and karyotypes of two hedgehogs (Insectivora: Erinaceidae) from Iran.

Karyological features of Erinaceus concolor and Hemiechinus auritus were studied from Zenjan (North Iran). The diploid number of chromosomes (2n), the total numbers of chromosomal arms (NF) and the numbers of autosomal arms (NFa) were determined as 2n = 48, NF = 92, NFa = 88 for E. concolor and as 2n = 50, NF = 98, NFa = 94 for H. auritus. Although the same dental formulae were established as 3.1.3.3/2.1.2.3 = 36 for both of species, they can be identified with some dental and cranial features from each other. Additionally, phallus of E. concolor was described.

Evolution of sensory specializations in insectivores.

Although insectivores have traditionally been thought of as primitive mammals with few specializations, recent studies have revealed great diversity in the sensory systems and brain organization of members of this mammalian order. The present article reviews some of these findings in three insectivore families that are thought to form a monophyletic group. These include hedgehogs (Erinaceidae), moles (Talpidae), and shrews (Soricidae). Members of each group live in unique ecological niches, have differently specialized senses, and exhibit different behaviors. Hedgehogs have well-developed visual, auditory, and somatosensory systems. Shrews make use of visual and auditory cues, but appear to depend most heavily on touch, particularly through prominent vibrissae. Moles are somatosensory specialists with small eyes and ears and unique epidermal mechanoreceptors called Eimer's organs used to identify prey and investigate their environment. In contrast to historical views of the insectivore order, members of this group have discrete and well-organized cortical sensory areas with sharp borders as determined from both electrophysiological mapping and analysis of cortical histology. Comparison of cortical organization across species reveals a number of specializations, including expansion of cortical representations of important sensory surfaces, the addition of cortical areas to some processing networks, and the subdivision of areas into separate cortical modules. In the case of the star-nosed mole, the somatosensory system has a tactile fovea and shares a number of features in common with the visual systems of sighted mammals.