Ancient DNA of narrow-headed vole reveal common features of the Late Pleistocene population dynamics in cold-adapted small mammals.

The narrow-headed vole, collared lemming and common vole were the most abundant small mammal species across the Eurasian Late Pleistocene steppe-tundra environment. Previous ancient DNA studies of the collared lemming and common vole have revealed dynamic population histories shaped by climatic fluctuations. To investigate the extent to which species with similar adaptations share common evolutionary histories, we generated a dataset comprised the mitochondrial genomes of 139 ancient and 6 modern narrow-headed voles from several sites across Europe and northwestern Asia covering approximately the last 100 thousand years (kyr). We inferred Bayesian time-aware phylogenies using 11 radiocarbon-dated samples to calibrate the molecular clock. Divergence of the main mtDNA lineages across the three species occurred during marine isotope stages (MIS) 7 and MIS 5, suggesting a common response of species adapted to open habitat during interglacials. We identified several time-structured mtDNA lineages in European narrow-headed vole, suggesting lineage turnover. The timing of some of these turnovers was synchronous across the three species, allowing us to identify the main drivers of the Late Pleistocene dynamics of steppe- and cold-adapted species.

Large-scale spatial patterns of small-mammal communities in the Mediterranean region revealed by Barn owl diet.

Due to mainly opportunistic hunting behaviour of Barn owl can be its diet composition used for assessing local structure of small-mammal community. We evaluated the structure of small-mammal communities in the Mediterranean region by analysing Barn owl diet using own pellets and literature data (85 localities comprising 182,343 prey individuals). Contrary to widely accepted macroecological theory, we found a latitudinal increase of small-mammal alpha diversity, a less distinct west-east increase and lower diversity on islands. The mean prey weight decreased with increasing latitude, while on islands it decreased with increasing island area. The mean prey weight on islands was further negatively affected by mean land modification by human and positively affected by its range. The diet diversity on islands was not affected either by island area or its distance from the mainland. Its composition largely conformed to the main pattern pronounced over whole the region: an unexpected homogeneity of small-mammal community structure. Despite high beta diversity and large between-sample variation in species composition, Crocidura (+ Suncus etruscus) and murids (Apodemus, Mus, Rattus, in marginal regions partly replaced by gerbillids, Meriones or Microtus) composed more than 90% of owl prey in 92% of samples. Peak abundances of these widespread species are associated with a dynamic mosaic of dense patches of sparse herb vegetation and evergreen sclerophyllous shrublands interspersing areas of human activity, the dominant habitat of the inner Mediterranean and richest food resource for foraging Barn owls. The respective small-mammal species can be looked upon as invasive elements accompanying large scale human colonization of the region since the Neolithic and replacing original island biota. Our study documented that desertification of the Mediterranean played an important role in shaping inverse latitudinal gradient in diversity of small-mammals that contradicts to widely accepted mecroecological theory.

Spatial networks differ when food supply changes: Foraging strategy of Egyptian fruit bats.

Animals are faced with a range of ecological constraints that shape their behavioural decisions. Habitat features that affect resource abundance will also have an impact, especially as regards spatial distribution, which will in turn affect associations between the animals. Here we utilised a network approach, using spatial and genetic data, to describe patterns in use of space (foraging sites) by free-ranging Egyptian fruit bats (Rousettus aegyptiacus) at the Dakhla Oasis in Egypt. We observed a decrease in home range size during spring, when food availability was lowest, which was reflected by differences in space sharing networks. Our data showed that when food was abundant, space sharing networks were less connected and more related individuals shared more foraging sites. In comparison, when food was scarce the bats had few possibilities to decide where and with whom to forage. Overall, both networks had high mean degree, suggesting communal knowledge of predictable food distribution.

Highly divergent lineage of narrow-headed vole from the Late Pleistocene Europe.

During the Late Pleistocene, narrow-headed voles (Lasiopodomys gregalis) inhabited Eurasia's vast territories, frequently becoming the dominant small mammal species among steppe-tundra communities. We investigated the relationship between this species' European and Asiatic populations by sequencing the mtDNA genomes of two extant specimens from Russia and 10 individuals from five Central European sites, dated to the post-LGM period. Phylogenetic analyses based on a large portion of mtDNA genomes highly supported the positioning of L. gregalis within Arvicolinae. The phylogeny based on mtDNA cytochrome b sequences revealed a deep divergence of European narrow-headed voles from Asiatic ones and their sister position against the extant L. gregalis and L. raddei. The divergence of the European lineage was estimated to a minimum 230 thousand years ago. This suggest, contrary to the current biogeographic hypotheses, that during the interglacial periods narrow-headed vole did not retreat from Europe but survived the unfavourable conditions within the refugial areas. Based on this result, we propose to establish a cryptic species status for the Late Pleistocene European narrow-headed vole and to name this taxon Lasiopodomys anglicus.

Carnivore distribution across habitats in a central-European landscape: a camera trap study.

Quantitative data on local variation in patterns of occurrence of common carnivore species, such as the red fox, European badger, or martens in central Europe are largely missing. We conducted a study focusing on carnivore ecology and distribution in a cultural landscape with the use of modern technology. We placed 73 automated infra-red camera traps into four different habitats differing in water availability and canopy cover (mixed forest, wetland, shrubby grassland and floodplain forest) in the Polabí region near Prague, Czech Republic. Each habitat was represented by three or four spatially isolated sites within which the camera traps were distributed. During the year of the study, we recorded nine carnivore species, including the non-native golden jackal. Habitats with the highest numbers of records pooled across all species were wetland (1279) and shrubby grassland (1014); fewer records were made in mixed (876) and floodplain forest (734). Habitat had a significant effect on the number of records of badger and marten, and a marginally significant effect on fox. In terms of seasonal dynamics, there were significant differences in the distribution of records among seasons in fox, marginally significant in least weasel, and the occurrence among seasons did not differ for badger and marten. In the summer, fox and marten were more active than expected by chance during the day, while the pattern was opposite in winter when they were more active during the night. Our findings on habitat preferences and circadian and seasonal activity provided the first quantitative data on patterns whose existence was assumed on the basis of conventional wisdom. Our study demonstrates the potential of a long-term monitoring approach based on infra-red camera traps. Generally, the rather frequent occurrence of recorded species indicates that most carnivore species are thriving in current central-European landscapes characterized by human-driven disturbances and urbanization.

Enamel apatite crystallinity significantly contributes to mammalian dental adaptations.

The monophyodont molar teeth, prismatic enamel and the complexity of enamel microarchitecture are regarded as essential dental apomorphies of mammals. As prominent background factors of feeding efficiency and individual longevity these characters are crucial components of mammalian adaptive dynamics. Little is known, however, to which degree these adaptations are influenced by the crystallographic properties of elementary hydroxyapatite crystallites, the only inorganic component of enamel. In a miniature pig where individual molars differ significantly in duration of their development and in enamel resistance to attrition stress, we found highly significant differences between the molars in the size of crystallites, amount of microstrain, crystallinity and in enamel stiffness and elasticity, all clearly scaled with the duration of tooth calcification. The same pattern was found also in red deer bearing different molar type. The results suggest that the prolongation of tooth development is associated with an increase of crystallinity, i.e. the atomic order of enamel hydroxyapatite, an obvious component of micromechanical property of mature enamel. This relation could contribute to prolongation of dental development, characteristic of mammals in general. The aspects of enamel crystallinity, omitted in previous studies on mammalian and vertebrate dental evolution, are to be taken in account in these topics.

Pre-oral gut contributes to facial structures in non-teleost fishes.

Despite the wide variety of adaptive modifications in the oral and facial regions of vertebrates, their early oropharyngeal development is considered strictly uniform. It involves sequential formation of the mouth and pharyngeal pouches, with ectoderm outlining the outer surface and endoderm the inner surface, as a rule. At the extreme anterior domain of vertebrate embryos, the ectoderm and endoderm directly juxtapose and initial development of this earliest ecto-endoderm interface, the primary mouth, typically involves ectodermal stomodeal invagination that limits the anterior expansion of the foregut endoderm. Here we present evidence that in embryos of extant non-teleost fishes, oral (stomodeal) formation is preceded by the development of prominent pre-oral gut diverticula (POGD) between the forebrain and roof of the forming mouth. Micro-computed tomography (micro-CT) imaging of bichir, sturgeon and gar embryos revealed that foregut outpocketing at the pre-oral domain begins even before the sequential formation of pharyngeal pouches. The presence of foregut-derived cells in the front of the mouth was further confirmed by in vivo experiments that allowed specific tracing of the early endodermal lining. We show that POGD in sturgeons contribute to the orofacial surface of their larvae, comprising oral teeth, lips, and sensory barbels. To our knowledge, this is the first thorough evidence for endodermal origin of external craniofacial structures in any vertebrate. In bichir and gar embryos, POGD form prominent cranial adhesive organs that are characteristic of the ancient bauplan of free-living chordate larvae. POGD hence seem arguably to be ancestral for all ray-finned fishes, and their topology, pharyngeal-like morphogenesis and gene expression suggest that they are evolutionarily related to the foregut-derived diverticula of early chordate and hemichordate embryos. The formation of POGD might thus represent an ancestral developmental module with deep deuterostome origins.

Late Pleistocene-Holocene paleobiogeography of the genus Apodemus in Central Europe.

Wood mice of the genus Apodemus are an essential component of small mammal communities throughout Europe. Molecular data suggest the postglacial colonization of current ranges from south European glacial refugia, different in particular species. Yet, details on the course of colonization and Holocene history of particular species are not available, partly because of a lack of reliable criteria for species identification in the fossil record. Using a sample of extant species, we analyzed variation patterns and between-species overlaps for a large set of metric and non-metric dental variables and established the criteria enabling the reliable species identification of fragmentary fossil material. The corresponding biometrical analyses were undertaken with fossil material of the genus (2528 items, 747 MNI) from 22 continuous sedimentary series in the Czech Republic and Slovakia, from LGM to Recent. In Central Europe, the genus is invariantly absent in LGM assemblages but regularly appears during the Late Vistulian. All the earliest records belong to A. flavicollis, the species clearly predominating in the fossil record until the Late Holocene. A. uralensis accompanied it in all regions until the late Boreal when disappeared from the fossil record (except for Pannonia). A few items identified as A. sylvaticus had already appeared in the early Holocene assemblages, first in the western part of the region, yet the regular appearance of the species is mostly in the post-Neolithic age. A. agrarius appeared sparsely from the Boreal with a maximum frequency during the post-Neolithic period. The results conform well to the picture suggested by molecular phylogeography but demonstrate considerable differences among particular species in dynamic of the range colonization. Further details concerning Holocene paleobiogeography of individual species in the medium latitude Europe are discussed.

Mammalian enamel maturation: Crystallographic changes prior to tooth eruption.

Using the distal molar of a minipig as a model, we studied changes in the microstructural characteristics of apatite crystallites during enamel maturation (16-23 months of postnatal age), and their effects upon the mechanical properties of the enamel coat. The slow rate of tooth development in a pig model enabled us to reveal essential heterochronies in particular components of the maturation process. The maturation changes began along the enamel-dentine junction (EDJ) of the trigonid, spreading subsequently to the outer layers of the enamel coat to appear at the surface zone with a 2-month delay. Correspondingly, at the distal part of the tooth the timing of maturation processes is delayed by 3-5 month compared to the mesial part of the tooth. The early stage of enamel maturation (16-20 months), when the enamel coat is composed almost exclusively of radial prismatic enamel, is characterized by a gradual increase in crystallite thickness (by a mean monthly increment of 3.8 nm); and an increase in the prism width and thickness of crystals composed of elementary crystallites. The late stage of maturation (the last two months prior to tooth eruption), marked with the rapid appearance of the interprismatic matrix (IPM) during which the crystals densely infill spaces between prisms, is characterized by an abrupt decrease in microstrain and abrupt changes in the micromechanical properties of the enamel: a rapid increase in its ability to resist long-term load and its considerable hardening. The results suggest that in terms of crystallization dynamics the processes characterizing the early and late stage of mammalian enamel maturation represent distinct entities. In regards to common features with enamel formation in the tribosphenic molar we argue that the separation of these processes could be a common apomorphy of mammalian amelogenetic dynamics in general.

Deeply torpid bats can change position without elevation of body temperature.

Because body temperature is tightly coupled to physiological function, hibernating animals entering deep torpor are typically immobile. We analysed thermal behaviour and locomotory activity of hibernating greater mouse-eared bats Myotis myotis and found two types of movement behaviour related to body temperature, i.e. movement at high fur temperature and at low fur temperatures (Tflow; <5°C). First Tflow movements appeared at the beginning of March and often occurred during long torpor bouts. In most cases, Tflow events represented slow displacements between clusters of bats. In several cases, however, departure or arrivals from and into clusters was also recorded without any elevation in body temperature. Distance travelled, flight duration and speed of locomotion during Tflow events was lower than in high fur temperature events. Such behaviour could allow bats to save energy long-term and prolong torpor bouts. Tflow movement in torpid bats significantly changes our understanding of basic hibernation principles and we strongly recommend further studies on the subject.

Circum-Mediterranean phylogeography of a bat coupled with past environmental niche modeling: A new paradigm for the recolonization of Europe?

The isolation of populations in the Iberian, Italian and Balkan peninsulas during the ice ages define four main paradigms that explain much of the known distribution of intraspecific genetic diversity in Europe. In this study we investigated the phylogeography of a wide-spread bat species, the bent-winged bat, Miniopterus schreibersii around the Mediterranean basin and in the Caucasus. Environmental Niche Modeling (ENM) analysis was applied to predict both the current distribution of the species and its distribution during the last glacial maximum (LGM). The combination of genetics and ENM results suggest that the populations of M. schreibersii in Europe, the Caucasus and Anatolia went extinct during the LGM, and the refugium for the species was a relatively small area to the east of the Levantine Sea, corresponding to the Mediterranean coasts of present-day Syria, Lebanon, Israel, and northeastern and northwestern Egypt. Subsequently the species first repopulated Anatolia, diversified there, and afterwards expanded into the Caucasus, continental Europe and North Africa after the end of the LGM. The fossil record in Iberia and the ENM results indicate continuous presence of Miniopterus in this peninsula that most probably was related to the Maghrebian lineage during the LGM, which did not persist afterwards. Using our results combined with similar findings in previous studies, we propose a new paradigm explaining the general distribution of genetic diversity in Europe involving the recolonization of the continent, with the main contribution from refugial populations in Anatolia and the Middle East. The study shows how genetics and ENM approaches can complement each other in providing a more detailed picture of intraspecific evolution.

Golden jackal (Canis aureus) in the Czech Republic: the first record of a live animal and its long-term persistence in the colonized habitat.

A golden jackal (Canis aureus) individual was recorded ~40 km east of Prague in the Czech Republic. It is the first record of a living golden jackal in the country; up to now several individuals have been recorded but all of them were either shot dead or killed by a vehicle. The observed animal was documented by camera traps set up for research of carnivore diversity in different habitats in the study area. It was first photographed on 19 June 2015, and in total there were 57 records made by 12 traps until 24 March 2016 when the animal was still present in the area. Forty-nine of the 57 records were made in a shrubby grassland over an area of ~100 ha, 39% of sightings were during the day and 61% in the night. There were two distinct peaks in the circadian activity of the animal, from 4 to 10 a.m., and from 6 p.m. to midnight. We also review the verified records of the golden jackal in the Czech Republic, some of which were only published in local hunting magazines. However, the observation reported in this paper represents the first evidence of a long-term occurrence in Europe of the same golden jackal individual, that persisted for at least nine months and over winter, northwest of Hungarian-Austrian border where the population has been known to reproduce.

Snezna jama (Slovenia): Interdisciplinary dating of cave sediments and implication for landscape evolution.

Caves are important markers of surface evolution, since they are, as a general rule, linked with ancient valley bottoms by their springs. However, caves can only be dated indirectly by means of the sediments they contain. If the sediment is older than common dating methods, one has to use multiple dating approaches in order to get meaningful results. U/Th dating, palaeomagnetic analysis of flowstone and sediment profiles, cosmogenic dating of quartz pebbles, and mammalian dating allowed a robust estimate of speleogenesis, sediment deposition, climatic change at the surface, and uplift history on the Periadriatic fault line during the Plio-Pleistocene. Our dates indicate that Snezna jama was formed in the (Upper) Miocene, received its sedimentary deposits during the Pliocene in a rather low-lying, hilly landscape, and became inactive due to uplift along the Periadriatic and Sava faults and climatic changes at the beginning of the Quaternary. Although it is only a single cave, the information contained within it makes it an important site of the Southern Alps.

Testing hypotheses of bat baculum function with 3D models derived from microCT.

The baculum (os penis) has been extensively studied as a taxon-specific character in bats and other mammals but its mechanical function is still unclear. There is a wide consensus in the literature that the baculum is probably a sexually selected character. Using a novel approach combining postmortem manipulation and three-dimensional (3D) imaging, we tested two functional hypotheses in the common noctule bat Nyctalus noctula, the common pipistrelle Pipistrellus pipistrellus, and Nathusius' pipistrelle Pipistrellus nathusii: (i) whether the baculum can protect the distal urethra and urethral opening from compression during erection and copulation; and (ii) whether the baculum and corpora cavernosa form a functional unit to support both the penile shaft and the more distal glans tip. In freshly dead or frozen and thawed bats, we compared flaccid penises with artificially 'erect' penises that were inflated with 10% formalin. Penises were stained with alcoholic iodine and imaged with a lab-based high-resolution x-ray microtomography system. Analysis of the 3D images enabled us to compare the changes in relative positions of the baculum, corpora cavernosa, urethra, and corpus spongiosum with one another between flaccid and 'erect' penises. Our results support both functional hypotheses, indicating that the baculum probably performs two different roles during erection. Our approach should prove valuable for comparing and testing the functions of different baculum morphologies in bats and other mammals. Moreover, we have validated an essential component of the groundwork necessary to extend this approach with finite element analysis for quantitative 3D biomechanical modeling of penis function.

The BAARA (Biological AutomAted RAdiotracking) system: a new approach in ecological field studies.

Radiotracking is an important and often the only possible method to explore specific habits and the behaviour of animals, but it has proven to be very demanding and time-consuming, especially when frequent positioning of a large group is required. Our aim was to address this issue by making the process partially automated, to mitigate the demands and related costs. This paper presents a novel automated tracking system that consists of a network of automated tracking stations deployed within the target area. Each station reads the signals from telemetry transmitters, estimates the bearing and distance of the tagged animals and records their position. The station is capable of tracking a theoretically unlimited number of transmitters on different frequency channels with the period of 5-15 seconds per single channel. An ordinary transmitter that fits within the supported frequency band might be used with BAARA (Biological AutomAted RAdiotracking); an extra option is the use of a custom-programmable transmitter with configurable operational parameters, such as the precise frequency channel or the transmission parameters. This new approach to a tracking system was tested for its applicability in a series of field and laboratory tests. BAARA has been tested within fieldwork explorations of Rousettus aegyptiacus during field trips to Dakhla oasis in Egypt. The results illustrate the novel perspective which automated radiotracking opens for the study of spatial behaviour, particularly in addressing topics in the domain of population ecology.

Late cenozoic history of the genus Micromys (mammalia, rodentia) in Central Europe.

Molecular phylogeography suggests that Micromys minutus, the sole extant species of the genus, colonized its extensive range quite recently, during the Late Pleistocene-Holocene period. Rich Pliocene and Pleistocene fossil records both from Europe and China suggest rather continuous and gradual in situ phenotype rearrangements from the Pliocene to the Recent periods. To elucidate the discrepancy we reexamined a considerable part of the European fossil record of the genus (14 sites from MN15 to Q3, 0.4-4.2 Ma, including the type series of M. preaminutus from MN15 Csarnóta 2), analyzed them with the aid of detailed morphometric comparisons, and concluded that: (a) The European Pliocene form, M. praeminutus, differs significantly from the extant species; (b) it exhibits a broad phenotypic variation covering the presumptive diagnostic characters of MN16 M. caesaris; (c) despite having smaller dimensions, the Early and Middle Pleistocene forms (MN17-Q3, 2.6-0.4 Ma) seem to be closer to M. praeminutus than to the extant species; (d) the extinction of M. praeminutus during Q3 and the re-occupation of its niche by the recent expansion of M. minutus from E-European-C Asiatic sources (suggested by phylogeographic hypotheses) cannot be excluded. Discussing interpretations of the phylogenetic past of the genus we emphasize the distinct history of the West Palearctic clade (Late Miocene-Early Pleistocene) terminating with M. praeminutus and the East Asiatic clade (chalceus, tedfordi, minutus), and the possible identity of the Western clade with the Late Miocene genus Parapodemus.

Development and evolution of the vertebrate primary mouth.

The vertebrate oral region represents a key interface between outer and inner environments, and its structural and functional design is among the limiting factors for survival of its owners. Both formation of the respective oral opening (primary mouth) and establishment of the food-processing apparatus (secondary mouth) require interplay between several embryonic tissues and complex embryonic rearrangements. Although many aspects of the secondary mouth formation, including development of the jaws, teeth or taste buds, are known in considerable detail, general knowledge about primary mouth formation is regrettably low. In this paper, primary mouth formation is reviewed from a comparative point of view in order to reveal its underestimated morphogenetic diversity among, and also within, particular vertebrate clades. In general, three main developmental modes were identified. The most common is characterized by primary mouth formation via a deeply invaginated ectodermal stomodeum and subsequent rupture of the bilaminar oral membrane. However, in salamander, lungfish and also in some frog species, the mouth develops alternatively via stomodeal collar formation contributed both by the ecto- and endoderm. In ray-finned fishes, on the other hand, the mouth forms via an ectoderm wedge and later horizontal detachment of the initially compressed oral epithelia with probably a mixed germ-layer derivation. A very intriguing situation can be seen in agnathan fishes: whereas lampreys develop their primary mouth in a manner similar to the most common gnathostome pattern, hagfishes seem to undergo a unique oropharyngeal morphogenesis when compared with other vertebrates. In discussing the early formative embryonic correlates of primary mouth formation likely to be responsible for evolutionary-developmental modifications of this area, we stress an essential role of four factors: first, positioning and amount of yolk tissue; closely related to, second, endoderm formation during gastrulation, which initiates the process and constrains possible evolutionary changes within this area; third, incipient structure of the stomodeal primordium at the anterior neural plate border, where the ectoderm component of the prospective primary mouth is formed; and fourth, the prime role of Pitx genes for establishment and later morphogenesis of oral region both in vertebrates and non-vertebrate chordates.

Tooth development in a model reptile: functional and null generation teeth in the gecko Paroedura picta.

This paper describes tooth development in a basal squamate, Paroedura picta. Due to its reproductive strategy, mode of development and position within the reptiles, this gecko represents an excellent model organism for the study of reptile development. Here we document the dental pattern and development of non-functional (null generation) and functional generations of teeth during embryonic development. Tooth development is followed from initiation to cytodifferentiation and ankylosis, as the tooth germs develop from bud, through cap to bell stages. The fate of the single generation of non-functional (null generation) teeth is shown to be variable, with some teeth being expelled from the oral cavity, while others are incorporated into the functional bone and teeth, or are absorbed. Fate appears to depend on the initiation site within the oral cavity, with the first null generation teeth forming before formation of the dental lamina. We show evidence for a stratum intermedium layer in the enamel epithelium of functional teeth and show that the bicuspid shape of the teeth is created by asymmetrical deposition of enamel, and not by folding of the inner dental epithelium as observed in mammals.

Increasing incidence of Geomyces destructans fungus in bats from the Czech Republic and Slovakia.

BACKGROUND: White-nose syndrome is a disease of hibernating insectivorous bats associated with the fungus Geomyces destructans. It first appeared in North America in 2006, where over a million bats died since then. In Europe, G. destructans was first identified in France in 2009. Its distribution, infection dynamics, and effects on hibernating bats in Europe are largely unknown. METHODOLOGY/PRINCIPAL FINDINGS: We screened hibernacula in the Czech Republic and Slovakia for the presence of the fungus during the winter seasons of 2008/2009 and 2009/2010. In winter 2009/2010, we found infected bats in 76 out of 98 surveyed sites, in which the majority had been previously negative. A photographic record of over 6000 hibernating bats, taken since 1994, revealed bats with fungal growths since 1995; however, the incidence of such bats increased in Myotis myotis from 2% in 2007 to 14% by 2010. Microscopic, cultivation and molecular genetic evaluations confirmed the identity of the recently sampled fungus as G. destructans, and demonstrated its continuous distribution in the studied area. At the end of the hibernation season we recorded pathologic changes in the skin of the affected bats, from which the fungus was isolated. We registered no mass mortality caused by the fungus, and the recorded population decline in the last two years of the most affected species, M. myotis, is within the population trend prediction interval. CONCLUSIONS/SIGNIFICANCE: G. destructans was found to be widespread in the Czech Republic and Slovakia, with an epizootic incidence in bats during the most recent years. Further development of the situation urgently requires a detailed pan-European monitoring scheme.

Enamel microarchitecture of a tribosphenic molar.

The tribosphenic molar is a dental apomorphy of mammals and the molar type from which all derived types originated. Its enamel coat is expected to be ancestral: a thin, evenly distributed layer of radial prismatic enamel. In the bat Myotis myotis, we reinvestigated the 3D architecture of the dental enamel using serial sectioning combined with scanning electron microscopy analyses, biometrics of enamel prisms and crystallites, and X-ray diffraction. We found distinct heterotopies in enamel thickness (thick enamel on the convex sides of the crests, thin on the concave ones), angularity of enamel prisms, and in distribution of particular enamel types (prismatic, interprismatic, aprismatic) and demonstrated structural relations of these heterotopies to the cusp and crest organization of the tribosphenic molar. X-ray diffraction demonstrated that the crystallites composing the enamel are actually the aggregates of much smaller primary crystallites. The differences among particular enamel types in degree of crystallite aggregation and the variation in structural microstrain of the primary crystallites (depending upon the duration and the mechanical context of mineralization) represent factors not fully understood as yet that may contribute to the complexity of enamel microarchitecture in a significant way.