Bone microstructure as an indicator of digging ability in moles (Talpidae, Eulipotyphla).

Talpid moles (Talpidae, Eulipotyphla) are mammals highly specialised in burrowing using their forelimbs. Fossoriality has allowed moles to expand their ecological niche by enabling access to subterranean resources and spaces. This specialisation in burrowing has led to adaptations in the forelimb bones of moles for humeral rotation digging, a distinctive strategy unparalleled among other diggers. While bone robustness has been examined in moles through external morphology, the adaptation of bone microstructure to digging strategy remains unclear. Based on two assumptions, (1) the humerus of moles is subjected to a torsional load due to humeral rotation digging, and (2) the magnitude of torsional load correlates with the compactness of the substrate in which the individuals can dig, we hypothesised that humeral rotation digging influences bone microstructure. Comparative analyses of transverse sections from the humeri and femora of three mole species (Mogera imaizumii, Mogera wogura and Urotrichus talpoides; Talpidae) and an outgroup eulipotyphlan (Suncus murinus; Soricidae) revealed that (1) vascular canals distributed in the humeri of moles align more predominantly circumferential along the bone walls, indicating an adaptation to the torsion generated by humeral rotation digging, and (2) the laminarity of vascular canals, particularly in Mogera species compared with Urotrichus, potentially reflects differences in the magnitude of load due to substrate compactness during digging. The aligned vascular canals are distinctive traits not observed in mammals employing other digging strategies. This suggests that vascular canal laminarity can be an indicator of not only humeral rotation digging in fossorial animals, but also the variation of eco-spaces in talpid species.

Burrowing star-nosed moles (Condylura cristata) are not hypoxia tolerant.

Star-nosed moles (Condylura cristata) have an impressive diving performance and burrowing lifestyle, yet no ventilatory data are available for this or any other talpid mole species. We predicted that, like many other semi-aquatic and fossorial small mammals, star-nosed moles would exhibit: (i) a blunted (i.e. delayed or reduced) hypoxic ventilatory response, (ii) a reduced metabolic rate and (iii) a lowered body temperature (Tb) in hypoxia. We thus non-invasively measured these variables from wild-caught star-nosed moles exposed to normoxia (21% O2) or acute graded hypoxia (21-6% O2). Surprisingly, star-nosed moles did not exhibit a blunted HVR or decreased Tb in hypoxia, and only manifested a significant, albeit small (<8%), depression of metabolic rate at 6% O2 relative to normoxic controls. Unlike small rodents inhabiting similar niches, star-nosed moles are thus intolerant to hypoxia, which may reflect an evolutionary trade-off favouring the extreme sensory biology of this unusual insectivore.

Karyotype Analysis of the New Talpa Species Talpa aquitania (Talpidae; Insectivora) from Northern Spain.

Karyotypes of 3 male Talpa specimens from northern Spain were analyzed. The mesostyles of upper molars and cytochrome b sequence analysis identified these specimens as belonging to Talpa aquitania, a new Talpa species recently described from northern Spain and southern France. We describe here for the first time the karyotype of Talpa aquitania. Its diploid number is 2n = 34 and NFa = 64, and all chromosomes including the sex chromosomes are biarmed, either metacentric or submetacentric. G-banding demonstrated that the karyotypes of T. aquitania and T. occidentalis (the most closely related species) are almost identical. However, the karyotype of T. aquitania differs from the karyotypes of both T. europaea and T. occidentalis in that it has a medium-sized biarmed Y chromosome rather than a dot-like chromosome and that chromosome 16 is submetacentric in T. aquitania but has a small p-arm in both T. europaea and T. occidentalis. Pericentromeric C-bands were scarce and only clearly visible in a few chromosomal pairs. In addition, C-banding demonstrated that half of the 14p, the 16p, and the Y chromosome are all heterochromatic. rDNA genes were located at the secondary constriction in autosomal pair 3, a common feature in the karyotypes of all Talpa species. Hybridization signals for telomeric repeats were found on the telomeres and the pericentric regions of some chromosomes and co-localized in the secondary constriction of pair 3 with the rDNA genes. In conclusion, the karyotype of T. aquitania from northern Spain is very similar to the karyotype of other species belonging to the genus Talpa.

Talpid Mole Phylogeny Unites Shrew Moles and Illuminates Overlooked Cryptic Species Diversity.

The mammalian family Talpidae (moles, shrew moles, desmans) is characterized by diverse ecomorphologies associated with terrestrial, semi-aquatic, semi-fossorial, fossorial, and aquatic-fossorial lifestyles. Prominent specializations involved with these different lifestyles, and the transitions between them, pose outstanding questions regarding the evolutionary history within the family, not only for living but also for fossil taxa. Here, we investigate the phylogenetic relationships, divergence times, and biogeographic history of the family using 19 nuclear and 2 mitochondrial genes (∼16 kb) from ∼60% of described species representing all 17 genera. Our phylogenetic analyses help settle classical questions in the evolution of moles, identify an ancient (mid-Miocene) split within the monotypic genus Scaptonyx, and indicate that talpid species richness may be nearly 30% higher than previously recognized. Our results also uniformly support the monophyly of long-tailed moles with the two shrew mole tribes and confirm that the Gansu mole is the sole living Asian member of an otherwise North American radiation. Finally, we provide evidence that aquatic specializations within the tribes Condylurini and Desmanini evolved along different morphological trajectories, though we were unable to statistically reject monophyly of the strictly fossorial tribes Talpini and Scalopini.

Spatio-temporal analysis of Nova virus, a divergent hantavirus circulating in the European mole in Belgium.

Over the last decade, the recognized host range of hantaviruses has expanded considerably with the discovery of distinct hantaviruses in shrews, moles and bats. Unfortunately, in-depth studies of these viruses have been limited. Here we describe a comprehensive analysis of the spatial distribution, genetic diversity and evolution of Nova virus, a hantavirus that has the European mole as its natural host. Our analysis demonstrated that Nova virus has a high prevalence and widespread distribution in Belgium. While Nova virus displayed relatively high nucleotide diversity in Belgium, amino acid changes were limited. The nucleocapsid protein was subjected to strong purifying selection, reflecting the strict evolutionary constraints placed upon Nova virus by its host. Spatio-temporal analysis using Bayesian evolutionary inference techniques demonstrated that Nova virus had efficiently spread in the European mole population in Belgium, forming two distinct clades, representing east and west of Belgium. The influence of landscape barriers, in the form of the main waterways, on the dispersal velocity of Nova virus was assessed using an analytical framework for comparing Bayesian viral phylogenies with environmental landscape data. We demonstrated that waterways did not act as an environmental resistance factor slowing down Nova virus diffusion in the mole population. With this study, we provide information about the spatial diffusion of Nova virus and contribute sequence information that can be applied in further functional studies.

Multilocus phylogeny of talpine moles (Talpini, Talpidae, Eulipotyphla) and its implications for systematics.

The tribe Talpini is a group of strictly subterranean moles distributed across the Eurasian Continent whose phylogenetic relationships and taxonomy remain unresolved. Here we report a multi-locus nuclear-mitochondrial DNA dataset (9468 bp) from 11 talpine species encompassing all five recognized genera, together with analyses of their divergence times and evolutionary affinities inferred from maximum likelihood and Bayesian approaches. Our results finely resolved all relationships except the root of the four recognized Asian genera, which was placed sister to the genus Talpa. With respect to the Asian clade, we moreover provide the first molecular support for a sister-taxon relationship between Parascaptor and Scaptochirus and confirm that the genus Euroscaptor is paraphyletic. Further, and despite a relatively small sample size (22 specimens), our species delimitation analyses support the existence of at least two genetically distinct, and hence potentially cryptic species. Taken together, these findings argue that generic status should be given to E. mizura and illustrate that the taxonomic diversity of the tribe Talpini in mountainous regions of southwestern China and Southeast Asia is underestimated. Finally, results of our divergence time analyses support a rapid radiation of the endemic Asian genera in the late-Miocene, which temporally corresponds with enhanced aridity and cooling arising from a significant uplift of the Himalayan-Tibetan plateau.

The complete mitochondrial genome of the long-tailed mole: Scaptonyx fusicaudus (Eulipotyphla, Talpidae).

The long-tailed mole (Scaptonyx fusicaudus) belongs to a monotypic genus within the family Talpidae. It is a small semi-fossorial mammal mostly distributed in south-western China. In this study, we obtained the complete mitochondrial genome of S. fusicaudus. The genome is a total of 16,602 bp in length, containing 13 protein-coding genes (PCGs), 22 transfer RNA genes (tRNA), 2 ribosomal RNA genes (rRNA) and 2 non-coding regions, with a base composition of 33.51% A, 28.73% T, 23.68% C and 14.08% G. The nucleotide sequence data of 13 protein-coding genes of S. fusicaudus and other 14 insectivora species were used to reconstruct a Bayesian phylogenetic tree. The tree shows that S. fusicaudus belongs to the subfamily Talpinae and is closely related to Urotrichus talpoides.

Comparison and Phylogenetic Analysis of Mitochondrial Genomes of Talpidae Animals.

Talpidae is a model group for evolutionary studies due to their highly specialized morphologies and diverse lifestyles. Mitochondrial genomes are molecular markers commonly used in species evolution and phylogenetic studies. In this study, the complete mitochondrial genome sequence of Scaptochirus moschatus was obtained by Illumina NovaSeq sequencing. The complete mitochondrial genomes of 14 Talpidae species (including Scaptochirus moschatus obtained in the present study) and the cytochrome b (Cyt b) gene sequences of 48 Talpidae species were downloaded from the NCBI database for comparison and phylogenetic studies to analyze the phylogenetic relationships and to find the possible reasons of the niche differentiation and ecotype specialization of Talpidae animals. The results showed that the mitochondrial genome sequences of 14 species belonging to the family Talpidae were 16,528 to 16,962 bp, all containing 13 protein-coding genes, 22 tRNA, two rRNA, and a non-coding region (control region). The difference in the number of repetitive repeats in the control region is responsible for the difference in the length of Talpidae mitochondrial genome sequences. Combining the divergence time of Talpidae animals with the geological history, it is found that the niche differentiation and ecotype divergence of Talpidae is closely related to historically global climate changes. Semi-aquatic groups diverged in the early Oligocene (about 31.22 MYA), probably in response to the global climate transition from warm to cool. During the early Miocene (about 19.54 MYA), some species of Talpidae moved to underground habitats and formed fossorial groups that were adept at digging due to the effects of the glaciation. In the middle Miocene (about 16.23 MYA), some Talpidae animals returned to the ground and formed semi-fossorial shrew moles as global climate warming again.

A NEW EIMERIAN (APICOMPLEXA: EIMERIIDAE) FROM EASTERN MOLE, SCALOPUS AQUATICUS (MAMMALIA: EULIPOTYPHLA: TALPIDAE), IN CENTRAL ARKANSAS, WITH ADDITIONAL INFORMATION ON CYCLOSPORA YATESI MCALLISTER, MOTRIUK-SMITH, AND KERR.

The eastern mole, Scalopus aquaticus (L.), is a common inhabitant of loamy soils in Canada, the eastern United States, and Mexico. Seven coccidian parasites have been previously reported from S. aquaticus, including 3 cyclosporans and 4 eimerians from hosts taken in Arkansas and Texas. A single S. aquaticus, collected in February 2022 in central Arkansas, was found to be passing oocysts of 2 coccidians, a new species of Eimeria, and Cyclospora yatesiMcAllister, Motriuk-Smith, and Kerr, 2018. Oocysts of Eimeria brotheri n. sp. are ellipsoidal (sometimes ovoidal) with a smooth bilayered wall, measure 14.0 × 9.9 µm, and have a length/width (L/W) ratio of 1.5; both micropyle and oocyst residua are absent, but a single polar granule is present. Sporocysts are ellipsoidal and measure 8.1 × 4.6 µm, L/W 1.8; a flattened to knoblike Stieda body as well as a rounded sub-Stieda body are present. The sporocyst residuum is composed of an irregular mass of large granules. Additional metrical and morphological information is provided on oocysts of C. yatesi. This study demonstrates that although several coccidians were previously documented from this host, additional S. aquaticus should be examined for coccidians from Arkansas as well as other parts of its range.

Mammals in Tawny Owl (Strixaluco) pellets from Kharkiv Region, Ukraine.

Background: The Tawny owl (Strixaluco) is a common owl species in Europe, demonstrating generalist diet strategy. Its main prey are small rodents and numerous studies show that the composition of its diet broadly reflects changes in prey species abundance in its habitats. Due to strictly sedentary habits of adult birds and their year-round territoriality, it is possible to locate habitats of their prey with a precision of several hundred metres. Analysis of owl pellets is a traditional method in faunistic studies to provide data on distribution of small mammals, especially cryptic species which are hard to be found using trapping. New information: Here, we present a dataset on mammals found in Tawny owl pellets collected during up to 13-year studies in the Kharkiv Region, Ukraine in three territories. Data from two territories were collected in a systematic way and allowed us to make analysis of seasonal, year-to-year and habitat variability in the Tawny owl diet and local mammal species composition.

Antimicrobial Activity of Cathelicidin-Derived Peptide from the Iberian Mole Talpa occidentalis.

The immune systems of all vertebrates contain cathelicidins, a family of antimicrobial peptides. Cathelicidins are a type of innate immune effector that have a number of biological functions, including a well-known direct antibacterial action and immunomodulatory function. In search of new templates for antimicrobial peptide discovery, we have identified and characterized the cathelicidin of the small mammal Talpa occidentalis. We describe the heterogeneity of cathelicidin in the order Eulipotyphla in relation to the Iberian mole and predict its antibacterial activity using bioinformatics tools. In an effort to correlate these findings, we derived the putative active peptide and performed in vitro hemolysis and antimicrobial activity assays, confirming that Iberian mole cathelicidins are antimicrobial. Our results showed that the Iberian mole putative peptide, named To-KL37 (KLFGKVGNLLQKGWQKIKNIGRRIKDFFRNIRPMQEA) has antibacterial and antifungal activity. Understanding the antimicrobial defense of insectivores may help scientists prevent the spread of pathogens to humans. We hope that this study can also provide new, effective antibacterial peptides for future drug development.

Satellitome Analysis on Talpa aquitania Genome and Inferences about the satDNAs Evolution on Some Talpidae.

In the genus Talpa a new species, named Talpa aquitania, has been recently described. Only cytogenetic data are available for the nuclear genome of this species. In this work, we characterize the satellitome of the T. aquitania genome that presents 16 different families, including telomeric sequences, and they represent 1.24% of the genome. The first satellite DNA family (TaquSat1-183) represents 0.558%, and six more abundant families, including TaquSat1-183, comprise 1.13%, while the remaining 11 sat-DNAs represent only 0.11%. The average A + T content of the SatDNA families was 50.43% and the median monomer length was 289.24 bp. The analysis of these SatDNAs indicated that they have different grades of clusterization, homogenization, and degeneration. Most of the satDNA families are present in the genomes of the other Talpa species analyzed, while in the genomes of other more distant species of Talpidae, only some of them are present, in accordance with the library hypothesis. Moreover, chromosomal localization by FISH revealed that some satDNAs are localized preferentially on centromeric and non-centromeric heterochromatin in T. aquitania and also in the sister species T. occidentalis karyotype. The differences observed between T. aquitania and the close relative T. occidentalis and T. europaea suggested that the satellitome is a very dynamic component of the genomes and that the satDNAs could be responsible for chromosomal differences between the species. Finally, in a broad context, these data contribute to the understanding of the evolution of satellitomes on mammals.

The complete mitochondrial genome of short-faced mole (Scaptochirus moschatus, Talpidae).

The short-faced mole (Scaptochirus moschatus) is a unique Chinese mammal that lives in burrows for life. In this study, we used Illumina NovaSeq sequencing to obtain the complete mitochondrial genome of the short-faced mole. The total length of the genome is 16,699 bp, containing 13 protein-coding genes, 22 transfer RNA genes (tRNA), 2 ribosomal RNA genes (rRNA), and 1 control region, with a base composition of 33.82% A, 26.89% T, 25.27% C, and 14.01% G. Phylogenetic analysis of the Talpidae by using complete mitochondrial genome sequences of 14 Talpidae species shows that short-faced mole is closely related to Parascaptor leucura.

Unique bone microanatomy reveals ancestry of subterranean specializations in mammals.

Acquiring a subterranean lifestyle entails a substantial shift for many aspects of terrestrial vertebrates' biology. Although this lifestyle is associated with multiple instances of convergent evolution, the relative success of some subterranean lineages largely remains unexplained. Here, we focus on the mammalian transitions to life underground, quantifying bone microanatomy through high-resolution X-ray tomography. The true moles stand out in this dataset. Examination of this family's bone histology reveals that the highly fossorial moles acquired a unique phenotype involving large amounts of compacted coarse cancellous bone. This phenotype exceeds the adaptive optimum seemingly shared by several other subterranean mammals and can be traced back to some of the first known members of the family. This remarkable microanatomy was acquired early in the history of the group and evolved faster than the gross morphology innovations of true moles' forelimb. This echoes the pattern described for other lifestyle transitions, such as the acquisition of bone mass specializations in secondarily aquatic tetrapods. Highly plastic traits-such as those pertaining to bone structure-are hence involved in the early stages of different types of lifestyle transitions.

The complete mitochondrial genome of white-tailed mole (Parascaptor leucura).

The white-tailed mole (Parascaptor leucura) belongs to genus Parascaptor, which is a monotypic genus distributed across Southwestern China, Assam (India), Bengal, and Northern Burma, and Laos. In this study, we obtained the complete mitochondrial genome of Parascaptor leucura. The genome is total 16,875 bp in length, containing 13 protein-coding genes, 22 transfer RNA genes (tRNA), two ribosomal RNA genes (rRNA), and two non-coding regions, with a base composition of 33.5% A, 26.4% T, 25.7% C, and 14.3% G. The nucleotide sequence data of 13 protein-coding genes of P. leucura and other nine Eulipotyphla species were used to reconstruct a Bayesian phylogenetic tree. The tree shows that P. leucura belongs to subfamily Talpinae and is closely related to Scaptochirus moschatus.

Myoglobin primary structure reveals multiple convergent transitions to semi-aquatic life in the world's smallest mammalian divers.

The speciose mammalian order Eulipotyphla (moles, shrews, hedgehogs, solenodons) combines an unusual diversity of semi-aquatic, semi-fossorial, and fossorial forms that arose from terrestrial forbearers. However, our understanding of the ecomorphological pathways leading to these lifestyles has been confounded by a fragmentary fossil record, unresolved phylogenetic relationships, and potential morphological convergence, calling for novel approaches. The net surface charge of the oxygen-storing muscle protein myoglobin (ZMb), which can be readily determined from its primary structure, provides an objective target to address this question due to mechanistic linkages with myoglobin concentration. Here, we generate a comprehensive 71 species molecular phylogeny that resolves previously intractable intra-family relationships and then ancestrally reconstruct ZMb evolution to identify ancient lifestyle transitions based on protein sequence alone. Our phylogenetically informed analyses confidently resolve fossorial habits having evolved twice in talpid moles and reveal five independent secondary aquatic transitions in the order housing the world's smallest endothermic divers.

The shrews, moles and hedgehogs that surround us all belong to the same large group of insect-eating mammals. While most members in this 'Eulipotyphla order' trot on land, some, like moles, have evolved to hunt their prey underground. A few species, such as the water shrews, have even ventured to adopt a semi-aquatic lifestyle, diving into ponds and streams to retrieve insects. These underwater foragers share unique challenges, burning a lot of energy and losing heat at a high rate while not being able to store much oxygen. It is still unclear how these semi-aquatic habits have come to be: the fossil record is fragmented and several species tend to display the same adaptations even though they have evolved separately. This makes it difficult to identify when and how many times the Eulipotyphla species started to inhabit water. The protein myoglobin, which gives muscles their red color, could help in this effort. This molecule helps muscles to capture oxygen from blood, a necessary step for cells to obtain energy. Penguins, seals and whales, which dive to get their food, often have much higher concentration of myoglobin so they can spend extended amount of time without having to surface for air. In addition, previous work has shown that eight groups of mammalian divers carry genetic changes that help newly synthetized myoglobin proteins to not stick to each other. This means that these animals can store more of the molecule in their muscles, increasing their oxygen intake and delivery. He et al. therefore speculated that all semi-aquatic Eulipotyphla species would carry genetic changes that made their myoglobin less likely to clump together; underground species, which also benefit from absorbing more oxygen, would display intermediate alterations. In addition, reconstructing the myoglobin sequences from the ancestors of living species would help to spot when the transition to aquatic life took place. A variety of approaches were harnessed to obtain myoglobin and other sequences from 55 eulipotyphlan mammals, which then were used to construct a strongly supported family tree for this group. The myoglobin results revealed that from terrestrial to subterranean to semi-aquatic species, genetic changes took place that would diminish the ability for the proteins to stick to each other. This pattern also showed that semi-aquatic lifestyles have independently evolved five separate times ­ twice in moles, three times in shrews. By retracing the evolutionary history of specific myoglobin properties, He et al. shed light on how one of the largest orders of mammals has come to be fantastically diverse.

Demodex crocidurae, a New Demodecid Mite (Acariformes: Prostigmata) Parasitizing the Lesser White-Toothed Shrew and a Redescription of Demodex talpae from European Mole with Data on Parasitism in Soricomorpha.

Only six parasitic species of Demodecidae mite have thus far been described from the Soricomorpha, these being associated with the common shrew Sorex araneus Linnaeus, 1758, and the Mediterranean water shrew Neomys anomalus Cabrera, 1907 (two species from each host), and with the lesser white-toothed shrew Crocidura suaveolens (Pallas, 1811) and the European mole Talpa europaea Linnaeus, 1758 (one from each host species). Presently, Demodex crocidurae, a new species, has been described from the territory of Poland for C. suaveolens; in order to confirm its validity, it was necessary to redescribe D. talpae Hirst, 1921, from T. europaea, a demodecid species first described by Hirst in 1921 from England and then noted only in Poland. Both species colonized the hairy skin of the body in their hosts, where no disease symptoms of infestation were observed. However, D. crocidurae showed higher infection parameters (prevalence 100%, mean intensity 11.7, intensity range 3-26 individuals) than those of D. talpae (30.0%, 4.7, 2.0-8.0), possibly due to different host biology.

All in the Family - Touch Versus Olfaction in Moles.

Here I review, compare, and contrast the neurobiology and behavior of the common, eastern mole (Scalopus aquaticus) and the star-nosed mole (Condylura cristata). These two species are part of the same family (Talpidae) and have similar body size and general morphology. But they differ in sensory specializations, complexity of neocortical organization, and behavior. The star-nosed mole has an elaborate mechanosensory organ-the star-consisting of 22 epidermal appendages (rays) covered with 25,000 touch domes called Eimer's organs. This densely innervated structure is represented in the neocortex in three different somatosensory maps, each visible in flattened neocortical sections as a series of 11 modules representing the 11 rays from the contralateral body. The 11th ray is greatly magnified in primary somatosensory cortex (S1). Behavioral studies show the star is moved in a saccadic manner and the 11th ray is a high-resolution tactile fovea, allowing star-nosed moles to forage on small prey with unprecedented speed and efficiency. In contrast, common mole noses lack Eimer's organs, their neocortex contains only two cortical maps of the nose, and they cannot localize small prey. Yet common moles have exceptional olfactory abilities, sniffing in stereo to rapidly localize discrete odor sources originating from larger prey. In addition, common moles are shown to track odorant trails laid down by moving prey. These results highlight the surprising abilities of species once thought to be simple, and the usefulness of diverse species in revealing general principles of brain organization and behavior. Anat Rec, 2019. © 2019 American Association for Anatomy.

Mercury, Lead and Cadmium Concentrations in Talpa occidentalis and in Their Digeneans of the Genus Ityogonimus.

INTRODUCTION: Many parasites living in aquatic ecosystems are useful indicators of environmental health. On the other hand, information is scarcer with respect to the use of helminth parasites of vertebrates living in terrestrial ecosystems as monitoring tools for toxic element environmental pollution. The present study evaluates the suitability of the model Talpa occidentalis/Ityogonimus spp. as a bioindicator system for mercury (Hg), lead (Pb) and cadmium (Cd) contamination in agricultural soils from Asturias (Spain). METHODS: Kidney and liver samples collected from T. occidentalis specimens (n = 36) and Ityogonimus spp. samples collected from 14 infected hosts were analyzed by ICP-MS. RESULTS: The highest mean levels of Hg and Pb were found in Ityogonimus individuals (20.9 and 12.4 µg g-1 wet weight, respectively). Considering renal and hepatic concentrations in T. occidentalis, bioaccumulation factors of Ityogonimus for Hg were 83.7 and 58.6, respectively, whereas concerning Pb bioaccumulation factors were 38.2 and 82.9, respectively. No bioaccumulation was detected in Ityogonimus in the case of Cd. CONCLUSIONS: More studies involving digenean parasites of small mammals are needed, especially when biomonitoring environmental toxic element pollution in terrestrial ecosystems. The present results support the above-mentioned model as a suitable biomonitoring system to evaluate environmental Hg and Pb contamination in terrestrial non-urban Iberian habitats. Similar models involving other species (Talpa spp./Ityogonimus spp.) might be used in a much wider geographical range.

Altered hemoglobin co-factor sensitivity does not underlie the evolution of derived fossorial specializations in the family Talpidae.

The high O2 affinity of European mole (Talpa europaea) blood is postulated to largely arise from the presence of two ß-globin chain residues (ß4 Ser and ß5 Gly) that weaken the interaction of its hemoglobin (Hb) with the red cell organophosphate 2,3-diphosphoglycerate (DPG). This latter trait is generally accepted to be an 'adaptation to subterranean life', despite the fact that no data are available for more basal mole lineages that have no evolutionary history of fossoriality (i.e. the ambulatory, high-elevation shrew-like moles and the semi-aquatic desmans), and which may similarly benefit from an elevated blood O2 affinity. To test whether evolution of a low DPG sensitivity phenotype is linked to derived fossorial lifestyles or represents an ancestral trait for the family, we determined the globin gene sequences and measured the intrinsic O2 affinity and co-factor sensitivity of the major Hb component of the gracile shrew-like mole (Uropsilus gracilis) and the Pyrenean desman (Galemys pyrenaicus). Our results unequivocally demonstrate that the presence of ß4 Ser and ß5 Gly, together with a low DPG sensitivity Hb phenotype, predates the radiation of the family Talpidae, and hence did not evolve as a specific adaptation to fossorial life. By contrast, our comparative analyses suggest that variations in whole blood O2 affinity among members of this family predominantly arose from amino acid substitutions that increase or decrease the intrinsic O2 affinity of the protein.