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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 unique reproductive and energetic features of lesser hedgehog tenrecs (Echinops telfairi) have been intensively studied in the field and within laboratory settings, but information on their propagation in zoological settings is limited. Based on a survey sent to zoological institutions currently housing reproductively active lesser hedgehog tenrecs in North America, this study reports on husbandry, veterinary practices, and demographics of reproducing lesser hedgehog tenrecs and their offspring. All 14 zoological institutions in the Lesser Hedgehog Tenrec Species Survival Plan who bred tenrecs within the last 12 years responded to the online survey. Provision of a nest box, nesting material, and a diet consisting of a commercial insectivore diet, insects, and produce was found in the majority of institutions, in addition to the practice of separating the breeding male and female before parturition. From the 24 dams representing 39 pregnancies and 158 young included in this study, the median calculated litter size was four young per litter. The median age of primiparity was 2.25 yr, and the oldest female to reproduce successfully was 10 yr old. The young had an overall rate of survival to weaning of 83%. A maternal complication rate of 26% and maternal mortality rate of 5% highlights the importance of veterinary care in periparturient dams, especially for detection and treatment of dystocia. These data provide guidance to zoological institutions breeding lesser hedgehog tenrecs.

Hibernation and daily torpor (heterothermy) allow endotherms to cope with demanding environmental conditions. The depth and duration of torpor bouts vary considerably between tropical and temperate climates, and tropical hibernators manage to cope with a wider spectrum of ambient temperature (Ta) regimes during heterothermy. As cycles in Ta can have profound effects on activity and torpor patterns as well as energy expenditure, we examined how these characteristics are affected by daily fluctuating versus constant Ta in a tropical hibernator, the lesser hedgehog tenrec (Echinops telfairi). Throughout the study, regardless of season, the tenrecs became torpid every day. In summer, E. telfairi used daily fluctuations in Ta to passively rewarm from daily torpor, which led to synchrony in the activity phases and torpor bouts between individuals and generally decreased energy expenditure. In contrast, animals housed at constant Ta showed considerable variation in timing and they had to invest more energy through endogenous heat production. During the hibernation season (winter) E. telfairi hibernated for several months in constant, as well as in fluctuating Ta and, as in summer, under fluctuating Ta arousals were much more uniform and showed less variation in timing compared to constant temperature regimes. The timing of torpor is not only important for its effective use, but synchronization of activity patterns could also be essential for social interactions, and successful foraging bouts. Our results highlight that Ta cycles can be an effective zeitgeber for activity and thermoregulatory rhythms throughout the year and that consideration should be given to the choice of temperature regime when studying heterothermy under laboratory conditions.

Causes of morbidity and mortality for various species of tenrecs have not been widely published, aside from several reports of neoplasia, and these data are crucial for advancing objectives for preventive medicine, diagnosis, and treatment. A survey on husbandry, morbidity, and mortality of lesser hedgehog tenrecs ( Echinops telfairi ) in Association of Zoos and Aquariums (AZA) institutions was conducted. Out of 32 institutions, 20 responded with data for 98 living and 93 dead animals. The most common causes of mortality among the dead animals were neoplasia (24%), hepatic lipidosis (11%), septicemia (8.6%), pneumonia (8.6%), cardiomyopathy (7.5%), renal disease (6.5%), osteomyelitis (3.2%), and trauma (3.2%). There was no statistically significant correlation between sex and neoplasia. Data about educational usage were specifically provided by survey respondents for 50 of the tenrecs, with only 42% being excluded from educational programming. Tenrecs are common to many AZA institutions as both educational and exhibit animals, and this study provides a helpful reference for expected health problems and highlights the need for future investments into medical diagnosis and treatment for these animals.

Currently, little is known about diseases and health concerns in tenrecs, family Tenrecidae. During the past 10 yr of breeding lesser Madagascar hedgehog tenrecs (Echinops telfairi) at a single zoological institution, a diverse range of obstetric and postpartum complications have been observed: pyometra, metritis, retained fetal membranes, retained fetus, perinatal mortality, and maternal neglect with subsequent successful hand-rearing of neonates. Treatment of these problems is complicated in tenrecs because of their small size and secretive behavior. Some of the cases reported in this article had successful resolution by ovariohysterectomy or cesarean section. The four cases described in this report overall help to disseminate options and outcomes for the diagnosis, treatment, and management of obstetric and postpartum complications in this small, secretive, nondomestic species.

1. Lesser Hedgehog Tenrec Echinops telfairi French: Petit Tenrec-hérisson / German: Kleiner Igeltenrek / Spanish: Tenrec erizo menor Other common names: Lesser Madagascar Tenrec Taxonomy. Echinops telfairi Martin, 1838, “Madagascar?” Echinops telfairi is considered to form a clade with Setifer setosus based on morphology and molecular studies. Monotypic. Distribution. W, SW & SE Madagascar, mainly S of Tsiribihina River. Descriptive notes. Head-body 104-165 mm, tail 13 mm, ear 16-25 mm, hindfoot 16-21 mm; weight 50-95 g. Crown of head and dorsal and lateral surfaces of body of the Lesser Hedgehog Tenrec are covered with dense, sharp, 10-12mm spines. Individuals vary from very pale to very dark; some spines can be dark over most of the length, light at tips, or dark for the entire length; no noticeable underfur is evident. Ventral surface, head, and limbs are covered with short, slightly bristly, light buffy hair. Ears are prominent. Dental formula is 2/2, C1/1,P 3/3, M 2/2 (x2) = 32. Habitat. Humid to sub-humid, dry, and dry spiny forests, open woodland, and degraded habitat from near sea level to elevations of 1300 m. The Lesser Hedgehog Tenrec occupies arboreal microhabitats. Food and Feeding. Captive Lesser Hedgehog Tenrecs eat species of Orthoptera and Annelida and occasionally frogs. Breeding. Lesser Hedgehog Tenrecs go through torpor for 3-5 months during cold season and begin mating when they emerge, usually in October. Gestation lasts 50-60 days, 61-64 days, or 62-68 days. Young are usually born in wet season when prey is most abundant. Litters have 1-6 young (4-5 embryos observed in wild specimens). Young are born nearly naked, with eyes and auditory meatus closed; eyes open at 7-9 days old; auditory meatus open at 14-16 days; and weaning begins at 18-23 days. Young begin to forage outside their nests with their mothers when they are c.3 weeks old, continuing to 4-5 weeks old. Activity patterns. The Lesser Hedgehog Tenrec is nocturnal, arboreal, and terrestrial. It shows daily and seasonal torpidity. Daily cycle includes gradual decline in body temperature and breathing rate to minima before noon, and they gradually increase to maxima at midnight. Lesser Hedgehog Tenrecs spend the day in nests of leaves and dead grasses in cavities oftree trunks and limbs, under logs, or at bases of trees, leaving the nest in early evening to forage. They enter torpor in winter. Movements, Home range and Social organization. Lesser Hedgehog Tenrecs are generally solitary when observed foraging in the wild, but they are apparently slightly socially tolerant; up to two individuals have been found together during seasonal torpor. Encounters in captivity between two males either led to mutual avoidance or fighting, which can be very aggressive. Scent marking occurs with deposition of feces and urine in specific places, and individuals drag their perineal regions across substrates when exploring new areas. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Lesser Hedgehog Tenrec presumably has a large overall, stable population and wide distribution. It occurs in protected areas and is tolerant of some habitat modification. Bibliography. Asher & Hofreiter (2006), Eibl-Eibesfeldt (1965), Eisenberg & Gould (1970), Everson et al. (2016), Goodman, Jenkins & Pidgeon (1999), Goodman, Soarimalala et al. (2013), Gould (1965), Gould & Eisenberg (1966), Herter (1962), Lovegrove & Génin (2008), Oelkrug et al. (2013), Olson & Goodman (2003), Poppitt et al. (1994), Poux, Madsen, Glos et al. (2008), Poux, Madsen, Marquard et al. (2005), Salton & Sargis (2008a, 2008b, 2009), Soarimalala & Goodman (2011), Stephenson, Racey & Rakotondraparany (1994), Stephenson, Soarimalala & Goodman (20164)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2018, Tenrecidae, pp. 134-172 in Handbook of the Mammals of the World – Volume 8 Insectivores, Sloths and Colugos, Barcelona :Lynx Edicions on page 163, DOI: 10.5281/zenodo.6808230

The lesser hedgehog tenrec (Echinops telfairi) displays reptile-like thermoregulatory behavior with markedly high variability in body temperature and metabolic rate. To understand how energy metabolism copes with this flexibility, we studied the bioenergetics of isolated liver mitochondria from cold (20°C) and warm (27°C) acclimated tenrecs. Different acclimation temperatures had no impact on mitochondrial respiration using succinate as the substrate. Mimicking the variation of body temperature by changing assay temperatures from 22 to 32°C highlighted temperature-sensitivity of respiration. The 40% reduction of respiratory control ratio (RCR) at 22°C compared to 32°C, a common estimate for mitochondrial efficiency, was caused by reduced substrate oxidation capacity. The simultaneous measurement of mitochondrial membrane potential enabled the precise assessment of efficiency with corrected respiration rates. Using this method, we show that proton leak respiration at the highest common membrane potential was not affected by acclimation temperature but was markedly decreased by assay temperature. Using membrane potential corrected respiration values, we show that the fraction of ATP-linked respiration (coupling efficiency) was maintained (70-85%) at lower temperatures. Collectively, we demonstrate that compromised substrate oxidation was temperature-compensated by the reduction of proton leak, thus maintaining the efficiency of mitochondrial energy conversion. Therefore, membrane potential data suggest that adjustments of mitochondrial proton leak contribute to energy homeostasis during thermoregulatory flexibility of tenrecs.

Studies of evolutionary developmental biology commonly use 'model organisms' such as fruit flies or mice, and questions are often functional or epigenetic. Phylogenetic investigations, in contrast, typically use species that are less common and mostly deal with broad scale analyses in the tree of life. However, important evolutionary transformations have taken place at all taxonomic levels, resulting in such diverse forms as elephants and shrews. To understand the mechanisms underlying morphological diversification, broader sampling and comparative approaches are paramount. Using a simple, standardized protocol, we describe for the first time the development of soft tissues and some parts of the skeleton, using μCT-imaging of developmental series of Echinops telfairi and Tenrec ecaudatus, two tenrecid afrotherian mammals. The developmental timing of soft tissue and skeletal characters described for the tenrecids is briefly compared with that of other mammals, including mouse, echidna, and the opossum. We found relatively few heterochronic differences in development in the armadillo vs. tenrec, consistent with a close relationship of Xenarthra and Afrotheria. Ossification in T. ecaudatus continues well into the second half of overall gestation, resembling the pattern seen in other small mammals and differing markedly from the advanced state of ossification evident early in the gestation of elephants, sheep, and humans.

A 15-yr-old, male lesser Madagascar hedgehog tenrec (Echinops telfairi) presented with a mass caudal to the right ear. Cytology suggested a sarcoma. Surgical removal was attempted. Histology was consistent with a soft tissue sarcoma. The mass recurred within 331 days post operation. Radiation therapy was initiated. Computed tomography was used for staging in conjunction with three-dimensional computerized treatment planning software to permit accurate lesion localization and to optimize normal tissue sparing. A total dose of 6,480 cGy was administered in 24 fractions over 46 days. Transient hind limb paresis developed during the course of the radiation therapy, but resolved after 7 days with prednisone treatment. Minimal acute radiation toxicity was observed. The mass responded with at least a 90% reduction in volume following radiation treatment. The animal survived 266 days from the initiation of treatment. On necropsy, a small mass and granulation tissue were found at the site of the initial neoplasm, indicating good regional control of the tumor; however, extensive metastases to the spleen and liver were present. Immunohistochemically, the original, recurrent, and metastatic populations were strongly positive for HMB 45 and weakly positive for S-100, and the final diagnosis was metastatic amelanotic melanoma.

Pituitary prolactin shows an episodic pattern of molecular evolution, with occasional short bursts of rapid change imposed on a generally rather slow evolutionary rate. In mammals, episodes of rapid change occurred in the evolution of primates, cetartiodactyls, rodents and the elephant. The bursts of rapid evolution in cetartiodactyls and rodents were followed by duplications of the prolactin gene that gave rise to large families of prolactin-related proteins including placental lactogens, while in primates the burst was followed by corresponding duplications of the related GH gene. The position in elephant is less clear. Extensive data relating to the genomic sequences of elephant and two additional members of the group Afrotheria are now available, and have been used here to characterize the prolactin genes in these species and explore whether additional prolactin-related genes are present. The results confirm the rapid evolution of elephant (Loxodonta africana) prolactin – the sequence of elephant prolactin is substantially different from that predicted for the ancestral placental mammal. Hyrax (Procavia capensis) prolactin is even more divergent but tenrec (Echinops telfairi) prolactin is strongly conserved. No evidence was obtained from searches of public databases for additional genes encoding prolactin-like proteins in any of these species. Detailed analysis of evolutionary rates, and other factors, indicates that the episode of rapid change in hyrax, and probably elephant, was adaptive, though the nature of the associated biological change(s) is not clear.

The patterns of heterothermy were measured in Lesser Hedgehog Tenrecs, Echinops telfairi, under semi-natural conditions in an outdoor enclosure during the austral mid-winter in southwestern Madagascar. The animals were implanted with miniaturized body temperature (Tb) loggers (iButtons) that measured body temperature every 42 min for 2 months (May and June). The tenrecs entered daily torpor on all 60 consecutive days of measurement, that is, on 100% of animal days, with body temperature closely tracking ambient temperature (Ta) during the ambient heating phase. The mean minimum daily Tb of the tenrecs was 18.44 +/- 0.50 degrees C (n = 174, N = 3), and never exceeded 25 degrees C whereas, apart from a few hibernation bouts in one animal, the mean maximum daily Tb was 30.73 +/- 0.15 degrees C (n = 167, N = 3). Thus during winter, tenrecs display the lowest normothermic Tb of all placental mammals. E. telfairi showed afternoon and early evening arousals, but entered torpor before midnight and remained in torpor for 12-18 h each day. One animal hibernated on two occasions for periods of 2-4 days. We consider E. telfairi to be a protoendotherm, and discuss the relevance and potential of these data for testing models on the evolution of endothermy.

It was purpose of this study to establish a safe and stable anaesthesia for the small hedgehog tenrec ( Echinops telfairi ) which can be used for short- and long-time interventions. Therefore 29 small hedgehog tenrecs were anaesthetized between 30 min and 4.5 h with a combination of the α 2 -agonist medetomidine (0.2 mg/kg), the benzodiazepine midazolam (3 mg/kg) and the dissociative anaesthetic ketamine (20 mg/kg) (MMK). All injections were administered subcutaneously (SC) in the area of the back by carefully lifting the animal's quills with a forceps. After SC injection of MMK animals lost their righting reflexes after 3.6 min (±1.12). Oxygen was supplemented to the animals’ nose and their body temperature was maintained constantly at 30 °C by a heating plate. Values of respiratory rate, pulse rate and oxygen saturation during the experiment were statistically evaluated by ANOVA and post-hoc tests to a level of significance determined as 5%. The animals had stable cardiovascular and respiratory values and good muscle relaxation. Between the 15th and the 45th minute the level of anaesthesia was deep enough for surgical interventions. Respiratory rate in this phase was 29.6±8.1 breaths/min and pulse rate was about 81.9±20 beats/min. MMK was partially antagonised with a combination of atipamezole (1 mg/kg) and flumazenile (0.2 mg/kg) (AF) SC. Time of complete recovery took about 8.8 min (8.76±4.31 min) after administering the antagonists. The partially antagonisable combination of MMK produced a stable anaesthesia in small hedgehog tenrecs up to 4.5 h. Therefore MMK can be used for short time interventions as well as e.g. for long-lasting neurophysiological recordings, when animals should survive the trials.

The otter shrews are members of the subfamily Potamogalinae within the family Tenrecidae. No description of the ovaries of any member of this subfamily has been published previously. The lesser hedgehog tenrec, Echinops telfairi, is a member of the subfamily Tenrecinae of the same family and, although its ovaries have not been described, other members of this subfamily have been shown to have ovaries with non-antral follicles. Examination of these two species illustrated that non-antral follicles were characteristic of the ovaries of both species, as was clefting and lobulation of the ovaries. Juvenile otter shrews range from those with only small follicles in the cortex to those with 300- to 400-µm follicles similar to those seen in non-pregnant and pregnant adults. As in other species, most of the growth of the oocyte occurred when follicles had one to two layers of granulosa cells. When larger follicles became atretic in the Nimba otter shrew, hypertrophy of the theca interna produced nodules of glandular interstitial tissue. In the tenrec, the hypertrophying theca interna cells in most large follicles appeared to undergo degeneration. Both species had some follicular fluid in the intercellular spaces between the more peripheral granulosa cells. It is suggested that this fluid could aid in separation of the cumulus from the remaining granulosa at ovulation. The protruding follicles in lobules and absence of a tunica albuginea might also facilitate ovulation of non-antral follicles. Ovaries with a thin-absent tunica albuginea and follicles with small-absent antra are widespread within both the Eulipotyphla and in the Afrosoricida, suggesting that such features may represent a primitive condition in ovarian development. Lobulated and deeply crypted ovaries are found in both groups but are not as common in the Eulipotyphla making inclusion of this feature as primitive more speculative.

Until recently, tenrecs were classified with insectivores in the order Lipotyphla, but nucleotide sequence data suggest they have closer affinities with a group of African mammals called Afrotheria. The placenta of Echinops has not been described and no studies involving electron microscopy of the placenta of any species of tenrec have been published. We used light and transmission electron microscopy to examine fixed placentae of embryos ranging from 25-66 mm in length. The placental disk is situated in the antimesometrial portion of the bicornuate uterus. The greater part of the disk consists of a labyrinth underlain by a spongy zone. The interhaemal barrier is unusual in that the trophoblastic component is a single layer of cytotrophoblast. These trophoblast cells have thick areas especially near the nuclei and extensive thin flanges but only occasionally have membrane-closed pore regions. The luminal surface has isolated patches of microvilli, and pinocytotic vesicles are numerous both apically and basally. In the centre of the placental disk is an elaborately folded haemophagous region. The primary folds have allantoic endoderm at one surface and columnar cytotrophoblast at the other. These trophoblast cells have numerous lipid droplets and vesicles, and often contain large yellow pigment crystalloids. The labyrinthine zone ends abruptly at the margins of the placental disk. However, the endoderm and connective tissue of the allantois and a layer of cytotrophoblast extend beyond the placental disk as a paraplacental region. Some of these distinctive features of Echinops placenta are shared with individual afrotherians, but no significant characteristic of definitive placentation is shared by all the Afrotheria.

The retinae of insectivores have been rarely studied, and their photoreceptor arrangements and expression patterns of visual pigments are largely unknown. We have determined the presence and distribution of cones in three species of shrews (common shrew Sorex araneus, greater white-toothed shrew Crocidura russula, dark forest shrew Crocidura poensis; Soricidae) and in the lesser hedgehog tenrec Echinops telfairi (Tenrecidae). Spectral cone types were identified and quantified in flattened whole retinae by antisera/antibodies recognizing the middle-to-long-wavelength-sensitive (M/L-)cone opsin and the short-wavelength-sensitive (S-)cone opsin, respectively. A combination of immunocytochemistry with conventional histology was used to assess rod densities and cone/rod ratios. In all four species the rods dominate at densities of about 230,000–260,000/mm2. M/L- and S-cones are present, comprising between 2% of the photoreceptors in the nocturnal Echinops telfairi and 13% in Sorex araneus that has equal diurnal and nocturnal activity phases. This suggests dichromatic color vision like in many other mammals. A striking feature in all four species are dramatically higher S-cone proportions in ventral than in dorsal retina (0.5% vs. 2.5–12% in Sorex, 5–15% vs. 30–45% in Crocidura poensis, 3–12% vs. 20–50% in Crocidura russula, 10–30% vs. 40–70% in Echinops). The functional and comparative aspects of these structural findings are discussed.

The phylogenetic position of the lesser hedgehog tenrec, Echinops telfairi, was studied on the basis of analysis of the concatenated sequences of 12 mitochondrial protein-coding genes. In addition to the tenrec, the analysis included two other representatives of the insectivore order Lipotyphla, the hedgehog and the mole. The eutherian tree was rooted with three non-eutherian mammalian taxa. The analysis joined the tenrec, the African elephant (order Proboscidea) and the aardvark (order Tubulidentata) on a common branch. The three lipotyphlan taxa, the tenrec (Tenrecidae), the mole (Talpidae) and the hedgehog (Erinaceidae) were dispersed in the eutherian tree, demonstrating lipotyphlan polyphyly.

To gain more insight into the evolution and functional significance of cerebrocerebellar circuits, the cerebellothalamic projections were studied with anterograde tracer substances in the Madagascan lesser hedgehog, tenrec. This insectivore shows one of the lowest size indices among mammals for both the cerebellar nuclei and the neocortex. Almost all cerebellodiencephalic target areas found in the tenrec have been described in other mammals. The intensity and extent of particular projections, however, vary considerably in the tenrec compared with the other mammals investigated so far. The most remarkable finding may be the tenrec's cerebellar projection to the nucleus ventralis medialis. This projection is the most prominent cerebellothalamic projection and originates in predominantly the lateral portion of the cerebellar nuclear complex. The projection to the caudolateral portion of the ventralis anterior complex (VAC) is located immediately rostral to the area receiving ascending somatosensory afferents and appears to originate, in particular, from the intermediate cerebellar nuclear complex. Another cerebellothalamic focus of terminations lies in the paralamellar region of the VAC, whereas the proper intralaminar nuclei, at best, receive a sparse cerebellar input. A faint-to-moderate projection, on the other hand, has been traced consistently to the ventral portion of the lateralis posterior-pulvinar complex and the adjacent dorsal geniculate nucleus. In addition, there are prominent cerebellosubthalamic projections to the zona incerta and the ventral geniculate nucleus. The latter projection is confined mainly to the ventralmost subdivision, which has been shown previously to receive ascending somatosensory, but not retinal, afferents. With the exception of the nucleus ventralis medialis, the projections were essentially confined to the contralateral side.

Endothermy has facilitated mammalian species radiation, but the sequence of events leading to sustained thermogenesis is debated in multiple evolutionary models. Here we study the Lesser hedgehog tenrec (Echinops telfairi), a phylogenetically ancient, ‘protoendothermic’ eutherian mammal, in which constantly high body temperatures are reported only during reproduction. Evidence for nonshivering thermogenesis is found in vivo during periodic ectothermic–endothermic transitions. Anatomical studies reveal large brown fat-like structures in the proximity of the reproductive organs, suggesting physiological significance for parental care. Biochemical analysis demonstrates high mitochondrial proton leak catalysed by an uncoupling protein 1 ortholog. Strikingly, bioenergetic profiling of tenrec uncoupling protein 1 reveals similar thermogenic potency as modern mouse uncoupling protein 1, despite the large phylogenetic distance. The discovery of functional brown adipose tissue in this ‘protoendothermic’ mammal links nonshivering thermogenesis directly to the roots of eutherian evolution, suggesting physiological importance prior to sustained body temperatures and migration to the cold., Endothermy facilitated mammalian species radiation, but the events leading to sustained thermogenesis are not clear. Here, the authors report functional brown adipose tissue in a protoendothermic mammal, linking nonshivering thermogenesis directly to the roots of eutherian endothermic evolution.

In a comparison of basic reproductive variables (number of teats, litter size, and duration of pregnancy) in the Madagascan Tenrecinae, some striking features (hyperthelia, litter size) are observed in Tenrec ecaudatus Lacépède, 1799 and the genus Hemicentetes Mivart, 1871. These variations are compared to Setifer setosus Froriep, 1806 and Echinops telfairi Martin, 1838. Tenrec ecaudatus gives birth to litters at intervals shorter than what was once believed to be the duration of pregnancy. This could be due either to superfetation, or to giving birth to one litter in two or more installments. The longitudinal stripe pattern of the genus Hemicentetes is interpreted as a mimicry of the sympatric, equally striped, juvenile Tenrec ecaudatus. The latter species protects its young with attending defensive-aggressive parents. The reproductive strategies of the Tenrecinae are discussed.

The aim of this study was to determine how a tropical tenrec, Echinops telfairi, is able to meet the obligatory extra costs of pregnancy and lactation. During pregnancy, mean body weight increased from 174.8 g (7.3 SD) to 279.0 g (10.0 SD), resting metabolic rate (RMR) from 28.7 kJ · d⁻¹ (0.4 SD) to 68.1 kJ · d⁻¹ (1.3 SD), and daily energy expenditure (DEE)from 42.1 kJ· d⁻¹ (3.5 SD) to 87.2 kJ · d⁻¹ (2.9 SD). Females actively maintained homeothermy throughout pregnancy. Food intake increased from 49.6 kJ · d⁻¹ (2.5 SD) to 121 kJ · d⁻¹ (13. 1 SD) during midpregnancy then gradually fell to 70.2 kJ · d⁻¹ (10. 6 SD) prior to parturition. Throughout lactation, body weight was stable at 262.6 g (4.3 SD). The mean RMR was 35.3 kJ · d⁻¹ (0. 8 SD), and DEE was 61.4 kJ · d⁻¹ (14.2 SD) when one pup was suckled. Homeothermy was also maintained throughout lactation. Food intake increased dramatically to 151 kJ · d⁻¹ (20.3 SD), and the energy transferred as milk to the litter was calculated to be between 34.9 kJ · d⁻¹ ...

The distribution of the neuropetide Y (NPY) was studied in geniculate and peri-chiasmatic regions in the lesser hedgehog-tenrec, Echinops telfairi (Insectivora). Only few neurons demonstrated NPY-like immunoreactivity in the ventral lateral geniculate nucleus. In contrast, NPY-immunoreactive perikarya were clearly present in the suprachiasmatic nucleus (SCh) and dorsal and caudal to it. The latter region might correspond to the subparaventricular zone (ZPV), recently identified in the rat as an additional area involved in processing circadian rhythms. While the distribution of a distinct cell population across nuclear boundares in both SCh and SPV might conform to the present idea of processing circadian rhythms, the presence of NPY-like immunoreactive neurons in these areas is rather unusual. In mammals, such neurons have only been demonstrated so far in the mentioned insectivore as well as in man.

The vomeronasal system (VNS) mediates pheromonal communication in mammals. From the vomeronasal organ, two populations of sensory neurons, expressing either Galphai2 or Galphao proteins, send projections that end in glomeruli distributed either at the rostral or caudal half of the accessory olfactory bulb (AOB), respectively. Neurons at the AOB contact glomeruli of a single subpopulation. The dichotomic segregation of AOB glomeruli has been described in opossums, rodents and rabbits, while Primates and Laurasiatheres present the Galphai2-pathway only, or none at all (such as apes, some bats and aquatic species). We studied the AOB of the Madagascan lesser tenrec Echinops telfairi (Afrotheria: Afrosoricida) and found that Galphai2 and Galphao proteins are expressed in rostral and caudal glomeruli, respectively. However, the segregation of vomeronasal glomeruli at the AOB is not exclusive, as both pathways contained some glomeruli transposed into the adjoining subdomain. Moreover, some glomeruli seem to contain intermingled afferences from both pathways. Both the transposition and heterogeneity of vomeronasal afferences are features, to our knowledge, never reported before. The organization of AOB glomeruli suggests that synaptic integration might occur at the glomerular layer. Whether intrinsic AOB neurons may make synaptic contact with axon terminals of both subpopulations is an interesting possibility that would expand our understanding about the integration of vomeronasal pathways.

Hemomonochorial placentation occurs in diverse species. We have examined placental glycosylation in five widely separated mammals with this type of placentation – lesser hedgehog tenrec (Echinops telfairi), spotted hyena (Crocuta crocuta), nine-banded armadillo (Dasypus novemcinctus), human (Homo sapiens) and guinea pig (Cavia porcellus) – in order to assess whether evolutionary convergence to the hemomonochorial state is accompanied by a similar convergence of glycan expression. Placentae from 2 E. telfairi, 3 C. crocuta, 1 D. novemcinctus, 4 womenand 1 C. porcellus were fixed and processed into epoxy resin. Binding of twenty-three lectins was assessed using a semiquantitative ranking system. The trophoblast apical/microvillous membrane of all five species showed marked similarities in glycosylation. In the N-linked series, there were abundant bi/tri-antennary complex chains, while the non-bisected variants were much scarcer. All species had plentiful N-acetyl lactosamine sequences; at chain termini, binding to Galβ1,4GlcNAc and Galβ1,3GalNAc sequences was greatly enhanced after neuraminidase treatment. In all species, terminal NeuNAcα2,3 residues were detected. The tenrec had unusually abundant terminal N-acetyl galactosamine. The basal plasma membrane/basal lamina showed glycosylation patterns distinct from the microvillous membrane in each case, indicating chemical diversity of the two opposite faces of trophoblast. Similar classes of glycan at the hemochorial interface suggest conservation of function. The observed lectin binding patterns suggest broad similarities of glycosylation that may have arisen by convergent evolution.

The gestation length (GL) of Tenrecs (Tenrecinae, Afrotheria) is still uncertain. This lack of knowledge also applies to the lesser hedgehog tenrec, Echinops telfairi, the species most commonly bred and maintained in captivity. The animals used in this study were held under controlled conditions (light, temperature and humidity). In order to determine the GL, groups of female tenrecs were subjected to various mating procedures followed by isolation periods of different lengths. A total of n = 249 pregnancies were analysed and the number of offspring per litter was 3.29 ± 0.09. The length of gestation could be determined in n = 199 pregnancies and a mean GL of 67.53 ± 0.36 days was calculated. Initial attempts with isolation periods of less than 16 days did not allow to accurately define the GL. Experiments with longer isolation periods and females subjected to only one mating procedure (n = 10) revealed a variation in the GLs of 57–79 days. However, in one female a GL of only 50 days was also observed indicating an even greater range in GL variation. There was a statistically significant tendency for shorter GLs in the animals that conceived later in the mating season, but no statistical evidence was found that age, parity or litter size played an essential role in determining the GL. In conclusion, an unexpected high variability in gestation length in E. telfairi was demonstrated although the study animals were kept under controlled environmental conditions. The factors and mechanisms regulating this high intra-species variability in gestation length need further investigations.

The hedgehog tenrec, Echinops telfairi, has certain basal mammalian features, like a cloaca and a sparsely differentiated brain with smooth cerebral hemispheres. The peripheral auditory capabilities of this species were investigated by means of distortion product otoacoustic emissions (DPOAE). For comparison, we determined auditory evoked potentials (AEP) in the inferior colliculus and the auditory cortex. Both methods show that the auditory range of E. telfairi extends well into ultrasonic frequencies, with a region of highest sensitivity at around 16 kHz. The total auditory range spans about 4 octaves at 40 dB SPL. The low-frequency limit of auditory processing is found at frequencies of about 2-3 kHz. The DPOAE and the AEP thresholds of E. telfairi do not run fully parallel in the high-frequency range. For a threshold value of 40 dB SPL, cochlear mechanical thresholds as measured with DPOAE extend up to 50 kHz, whereas neuronal thresholds reach the high-frequency limit at about 30 kHz. Frequency tuning, as assessed from DPOAE suppression tuning curves, was low to moderate with Q(10 dB) values ranging from 1.7 to 8. The lack of discontinuity in the group delay (derived from DPOAE measurements) reveals that cochlear frequency representation is tonotopic without any region of specialized mechanical tuning.

The mammalian cerebellum is histologically uniform. However, underlying the simple laminar architecture is a complex arrangement of parasagittal stripes and transverse zones that can be revealed by the expression of zebrin II/aldolase C. The cerebellar cortex of rodents, for example, is organized into four transverse zones: anterior, central, posterior and nodular. Within the anterior and posterior zones, parasagittal stripes of Purkinje cells expressing zebrin II alternate with those that do not. Zonal boundaries appear to be independent of cerebellar lobulation. To explore this model further, and to broaden our understanding of the evolution of cerebellar patterning, zebrin II expression has been studied in the cerebellum of the Madagascan hedgehog tenrec (Echinops telfairi), a basal insectivore with a lissiform cerebellum with only five lobules. Zebrin II expression in the tenrec reveals an array of four transverse zones as in rodents, two with homogeneous zebrin II expression, two further subdivided into stripes, that closely resembles the expression pattern described in other mammals. We conclude that a zone-and-stripe organization may be a common feature of the mammalian cerebellar vermis and hemispheres, and that zonal boundaries and cerebellar lobules and fissures form independently.

In order to find correlations between skin gland morphology and specific ethological features, the cutaneous glands of the foot pads of the primitive mammal the Madagascan tenrec, Echinops telfairi, were studied by histological and various histochemical methods as well as by electron microscopy. In the foot pads specific eccrine skin glands occurred consisting of coiled ducts and tubular secretory portions, the lumina of which were considerably wider than in primate sweat glands. The secretory tubules were composed of branched myoepithelial cells and glandular cells. The latter contained abundant mitochondria, large amounts of glycogen particles and few secretory granules as well as individual heterolysosomes and myelin bodies. The lateral cell membrane was marked by extensive interdigitations. The apical membranes of all glandular cells contained proteoglycans with sulfated and carboxylated groups containing N-acetyl-glucosamine, N-acetyl-galactosamine, galactose and mannose. The expression pattern of cytokeratins of the glandular epithelium was variable and showed similarities to that of the human eccrine glands. Tubulin, vinculin and actin were expressed in the glandular epithelium. The secretory cells showed positive reactions with antibodies against antimicrobial peptides and IgA. A positive reaction was observed with antibodies against the androgen receptor. The PCNA and TUNEL reactions indicated that the tubular skin glands of Echinops are made up of a slowly renewing tissue. We conclude that the glands fulfill several functions: production of a fluid-rich secretory product, which may prevent slipping of the foot pads on the substrate during running or climbing, secretion of antimicrobial peptides and proteins, and playing a role in thermoregulation.

The complete mitochondrial genome of a lesser hedgehog tenrec Echinops telfairi was determined in this study. It is an endemic African insectivore that is found specifically in Madagascar. The tenrec's back is covered with hedgehog-like spines. Unlike other spiny mammals, such as spiny mice, spiny rats, spiny dormice and porcupines, lesser hedgehog tenrecs look amazingly like true hedgehogs (Erinaceidae). However, they are distinguished morphologically from hedgehogs by the absence of a jugal bone. We determined the complete sequence of the mitochondrial genome of a lesser hedgehog tenrec and analyzed the results phylogenetically to determine the relationships between the tenrec and other insectivores (moles, shrews and hedgehogs), as well as the relationships between the tenrec and endemic African mammals, classified as Afrotheria, that have recently been shown by molecular analysis to be close relatives of the tenrec. Our data confirmed the afrotherian status of the tenrec, and no direct relation was recovered between the tenrec and the hedgehog. Comparing our data with those of others, we found that within-species variations in the mitochondrial DNA of lesser hedgehog tenrecs appear to be the largest recognized to date among mammals, apart from orangutans, which might be interesting from the view point of evolutionary history of tenrecs on Madagascar.

The rhinal cortex was investigated in the Madagascan lesser hedgehog tenrec, a basal placental mammal. This region parallels the rhinal indentation and presumably contains the equivalents of the insular and perirhinal cortices. Using cyto- and myeloarchitectural, enzyme- and immunohistochemical criteria as well as data on the connections with the olfactory bulb, the rhinal cortex was subdivided tentatively along its rostrocaudal and dorsoventral planes. An area caudally adjacent to the rhinal cortex received a prominent input from the olfactory bulb and was also preliminarily characterized in this study. Because previous studies in insectivores remained controversial with regard to the identification of the claustrum, special attention was paid to the laminar organization of the rhinal cortex and its deep cell groups. The tenrec’s claustrum was identified and delineated cytoarchitecturally and by its negative acetylcholinesterase stain. Latexin, a molecular marker for characterizing infragranular and claustral cells, also helped to differentiate the claustrum from the cell groups subjacent to it. Thus, the data indicate that in poorly differentiated mammals the claustrum occupies an intermediate deep position within the width of the rhinal cortex, i.e., it is separated from the subcortical white matter by additional, still unidentified, cell groups.

In order first to overcome the difficulties in understanding the increasing amount of information available regarding the mammalian somatosensory thalamus, and then to correlate the findings among different species and integrate them into a general concept of thalamic organization, the present study investigated the spinothalamic and medial lemniscal projections in Madagascan hedgehog tenrecs (Echinops telfairi and Setifer setosus). Tracer substances were injected into the dorsal column nuclei and into spinal segments at various levels; additional injections were made into the inferior colliculus. The ascending somesthetic projections were to predominantly contralateral posterolateral target areas, and were almost mirror-like on both sides to intralaminar and medial thalamic nuclei. The densest and most extensive projections, originating mainly from the high cervical spinal cord and the dorsal column nuclei, reached the posterolateral thalamus caudal to the lateral geniculate nucleus. This region was difficult to subdivide cytoarchitecturally; nevertheless, on the basis of its labeling pattern, several subdivisions could be described and preliminary named. Some of them compared tentatively with the internal portion of the medial geniculate nucleus (GM) and the ventral posterior nuclear complex (VPC) in more differentiated mammals. The most prominent subdivision, however, located subjacent to the lateral surface of the brainstem, was shown to receive additional fibers from the inferior colliculus. This region might be considered a further subdivision of GM, VPC, a perigeniculate area, and/or a region of its own not comparable at present, with thalamic regions in other mammals. On the other hand, it may also be a remnant of the hypothetical, diffuse multimodal region from which GM and VPC have possibly evolved.(ABSTRACT TRUNCATED AT 250 WORDS)

Echinops telfairi Martin, 1838. Proc. Zool. Soc. Lond., 1838:17. TYPE LOCALITY: Madagascar. DISTRIBUTION: S Madagascar. SYNONYMS: militarti, nigrescens, pallens., Published as part of Rainer Hutterer, 1993, Order Insectivora, pp. 69-130 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 73, DOI: 10.5281/zenodo.7353085

Using retrograde axonal flow and wheatgerm agglutinin conjugated to horseradish peroxidase, we studied the distribution of cortical neurons giving rise to spinal and dorsal column nuclear projections, and correlated the regions involved in the projections with the cytoarchitectonic areas recently identified in the lesser hedgehog tenrec, Echinops telfairi (Insectivora). Labeled cortical neurons were most numerous following injections of tracer into higher cervical segments, whereas almost none were found following thoracic injections. The cortical labeling appeared more prominent ipsilaterally than contralaterally after spinal injections, although it was more prominent on the contralateral side after injection into the dorsal column nuclear complex. The majority of labeled neurons found in lamina V occupied the neocortex adjacent to the interhemispheric fissure along the rostrocaudal extent of the small corpus callosum. This location corresponded to an intermediate rostrocaudal portion of the hemisphere, and particularly to area 2 of Rehkämper. In some cases, adjacent portions of areas 1 and 3 were also involved, as well as neocortical regions of the lateral hemisphere. The present data did not suggest a somatotopic organization of the projections; likewise, evidence for the presence of more than one somatosensorimotor representation was sparse.

Summary: The primary structures of the a- and /3hemoglobin chains of the lesser hedgehog tenrec (Echinops telfairi, Zalambdodonta) are presented. Chain separation was performed by carboxymethyl­ cellulose chromatography. The peptides, obtained by tryptic digestion of the oxidized chains, were prefrac­ tionated by gel chromatography and isolated by re­ versed·phase HPLC. For sequence analysis gas and liquid phase sequencers were employed. The tenrec hemoglobin consists of one a- and two /3-chains the latter occurring in a 1: 1 ratio and differing in /316 Glyl Cys and /3118 PhelLeu. Two external cysteine residues at /316 and /352 cause reversible polymerization to oc­ tamers and most likely irreversible formation of higher polymers. A comparison of the whole chains and certain posi­ tions of tenrec hemoglobin with those of Insectivora sensu strictu, Scandentia and Proto- and Metatheria corroborates a long and independent evolution of ten­ rec and its phylogenetic isolation from the Insectivora s.str. (hedgehog, musk shrew and mole). Replacements at positions involved in heme and sub­ unit interface contacts are discussed. Compared to human hemoglobin the temec pigment shows a low intrinsic oxygen affinity as well as lower chloride and temperature sensitivities, a reduced Bohr effect and a strong response to 2,3-DPG. The possible adaptive significance of these properties is discussed in relation to the large diurnal body temperature variations seen in tenrecs. Primiirstruktur und Sauerstoffbindungs-Eigenschaften des Hiimoglobins vom Kleinen Igeltanrek (Echinops telfairi, Zalambdodonta) - Ein Hinweis auf phylogenetische Isolation Zusammenfassnng: Die Primarstruktur der a- und /3Ketten des Hamoglobins vom Kleinen Igeltanrek (Echinops telfairi, Zalambdodonta) wird angegeben. Die Ketten konnten durch Carboxymethylcellulose­ Chromatographie getrennt werden. Die nach der tryptischen Spaltung der oxidierten Ketten erhalte­ nen Peptide wurden zunachst durch Gelchromatogra­ phie vorfraktioniert und dann mit Hilfe von Rever­ sed-Phase-HPLC isoliert. Die Sequenzanalyse er­ folgte in Gas- und Fltissigphasen-Sequenatoren. Das Tanrek-Hamoglobin hat eine a- und zwei /3-Ketten.

Using the autoradiographic tracing technique the retinal projections were studied in the tenrecs, Echinops telfairi and Setifer setosus (insectivora, tenrecidae). Bilateral projections were found to the n. suprachiasmaticus, the anterior hypothalamic area, the dorsal and ventral lateral geniculate bodies, the pretectal olivary nucleus and the superior colliculus. The contralateral projections were usually more intense than the ipsilateral ones except the retinohypothalamic connections. A partial segregation of the projection fields from both eyes was present in the dorsal and ventral lateral geniculate bodies. In the superior colliculus retinal fibers predominantly involved the stratum zonale and the upper portion of the stratum griseum superficiale on both sides. The projections to the deeper portion of the colliculi were rather faint, particularly on the ipsilateral side. Target areas receiving contralateral projections exclusively were the periamygdaloid area (labeled only in Setifer), the terminal accessory nuclei including the n. tractus optici and the inferior colliculus. The data are compared with other species. The most striking finding may concern the projection to the medial terminal nucleus being quite prominent in marsupials and most eutherian mammals (including the erinaceomorphous hedgehogs), but greatly reduced in tenrecs and primates.

1. Als neue Helminthenarten wurden bei Echinops telfairi (Centetidae: Insectivora) im Magen Physaloptera sprehni n. sp. und im Dunndarm Heteracanthorhynchus echinopsi n. sp. bestimmt.

Laboratory and field observations provide data on the life history and growth of Hemicentetes, Centetes, Echinops and Setifer . Comments on Geogale, Microgale ( Nesogale ) and Limnogale are also included. Similarities in courtship and agonistic behavior are noted. Some data on torpor indicate the irregularity of its occurrence, particularly in Hemicentetes semispinosus . A comparison of age at which the eyes open in four insectivore families indicates that tenrecs mature earliest, about 14 to 22 days in Talpidae, Soricidae and Erinaceidae, 7 to 13 days in Tenrecidae. Hemicentetes semispinosus and Echinops telfairi attain adult size at 6 to 8 weeks of age under laboratory conditions.

The Argas subgenus Secretargas Hoogstraal has previously been based only on A. (S.) transgariepinus White, a parasite of bats and geckos ranging from South Africa to southern Europe and the Near East. Discovery in Madagascar of adults and nymphs of A. hoogstraali Morel and Vassiliades, which was described from larvae only, provides information showing that this species also belongs to the subgenus Secretargas . Both sexes and large and small nymphal instars are described and the larva is redescribed. A. hoogstraali parasitizes 3 species of Oplurus lizards that shelter under stones on slopes that are flooded during the rainy season in the hot, semiarid southwestern area of Madagascar. A related parasite, A. (S.) echinops , new species, is also described from both sexes, large and small nymphs, and larvae. This species infests the Small Madagascar Hedgehog, Echinops telfairi Martin, and is found under larger stones in the same hot, semiarid savannah zone of Madagascar as A. (S.) hoogstraali .

This paper reports on possible effects of selective logging on vegetation structure and its consequences for two tenrec species in a dry forest in western Madagascar. On a small scale semi-mechanized logging of less than 10 m3 per ha alters forest structure significantly. However on a large scale most of these changes are smaller than the variation of forest structure due to natural causes. The most profound ecological effect of logging is a reduction in the number of woody species in the regenerating cohort of trees. Potential longterm consequences of this feature are discussed. Microhabitat variables influencing the habitat utilization of two tenrec species,Echinops telfairi andTenrec ecaudatus, were identified in an area of unlogged forest. The conclusions derived from this analysis were then tested in other parts of the forest which had been modified by selective logging. Logging of the extent described above slightly impairs habitats forE. telfairi but has no effect onT. ecaudatus. Though the latter species is hunted extensively in the logged areas, loggingper se does not threaten the survival of either species.

Echinops telfairi Martin, 1838. Proc. Zool. Soc. Lond., 1838:17. TYPE LOCALITY: Madagascar. DISTRIBUTION: S. Madagascar. ISIS NUMBER: 5301403002002001001., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Insectivora, pp. 58-106 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 58, DOI: 10.5281/zenodo.7353001