Kevin Campbell | University of Manitoba (original) (raw)

Papers by Kevin Campbell

Journal of Experimental Biology, 2000

The postprandial increase in metabolic rate associated with consuming, assimilating and excreting... more The postprandial increase in metabolic rate associated with consuming, assimilating and excreting a meal is often termed the heat increment of feeding (HIF). The metabolic heat production of star-nosed moles, Condylura cristata, held at thermoneutrality was monitored for 4 h following a single 10 min session of feeding on a ration consisting of 0 g (controls), 3.5 g or 10 g of earthworms. Coefficients for metabolizable energy digestibility and digesta passage rate of earthworms fed to C. cristata were also determined. We then tested whether feeding-induced thermogenesis substitutes partially or completely for thermoregulatory heat production in these animals exposed to sub-thermoneutral air temperatures (9-24 degrees C). A single feeding on earthworms had both short- and long-term effects on the metabolic rate and respiratory exchange ratio of C. cristata. The observed short-term (0-65 min) rise in metabolic rate, assumed to be associated primarily with the physical costs of nutrien...

ABSTRACTAllometry predicts that the 12–17 g American water shrew (Sorex palustris)—the world’s sm... more ABSTRACTAllometry predicts that the 12–17 g American water shrew (Sorex palustris)—the world’s smallest mammalian diver—will have the highest diving metabolic rate coupled with the lowest total body oxygen storage capacity, skeletal muscle buffering capacity, and glycolytic potential of any endothermic diver. Consistent with expectations, and potentially owing to their low thermal inertia, water shrews had a significantly higher diving metabolic rate in 10°C (8.77 mL O2 g−1 hr−1) compared to 30°C water (6.57 mL O2 g−1 hr−1). Unlike larger-bodied divers, muscle myoglobin contributed minimally (7.7–12.4%) to total onboard O2 stores of juvenile and adult water shrews, respectively, but was offset by high blood O2 carrying capacities (26.4–26.9 vol. %). Diving was predominantly aerobic, as only 2.3–3.9% of dives in 30 and 10°C water, respectively, exceeded the calculated aerobic dive limits at these temperatures (10.8–14.4 sec). The mean voluntary dive time of water shrews during 20-min...

The δ-globin gene (HBD) of eutherian mammals exhibits a propensity for recombinational exchange w... more The δ-globin gene (HBD) of eutherian mammals exhibits a propensity for recombinational exchange with the closely linked β-globin gene (HBB) and has been independently converted by the HBB gene in multiple lineages. Here we report the presence of a chimeric β/δ fusion gene in the African elephant (Loxodonta africana) that was created by unequal crossing-over between misaligned HBD and HBB paralogs. The recombinant chromosome that harbors the β/δ fusion gene in elephants is structurally similar to the ‘‘anti-Lepore’’ duplication mutant of humans (the reciprocal exchange product of the hemoglobin Lepore deletion mutant). However, the situation in the African elephant is unique in that the chimeric β/δ fusion gene supplanted the parental HBB gene and is therefore solely responsible for synthesizing the β-chain subunits of adult hemoglobin. A phylogenetic survey of β-like globin genes in afrotherian and xenarthran mammals revealed that the origin of the chimeric β/δ fusion gene and the c...

bioRxiv, 2021

Allometry predicts that the 12–17 g American water shrew (Sorex palustris)—the world’s smallest m... more Allometry predicts that the 12–17 g American water shrew (Sorex palustris)—the world’s smallest mammalian diver—will have the highest diving metabolic rate coupled with the lowest total body oxygen storage capacity, skeletal muscle buffering capacity, and glycolytic potential of any endothermic diver. Consistent with expectations, and potentially owing to their low thermal inertia, water shrews had a significantly higher diving metabolic rate in 10°C (8.77 mL O2 g−1 hr−1) compared to 30°C water (6.57 mL O2 g−1 hr−1). Unlike larger-bodied divers, muscle myoglobin contributed minimally (7.7–12.4%) to total onboard O2 stores of juvenile and adult water shrews, respectively, but was offset by high blood O2 carrying capacities (26.4–26.9 vol. %). Diving was predominantly aerobic, as only 2.3–3.9% of dives in 30 and 10°C water, respectively, exceeded the calculated aerobic dive limits at these temperatures (10.8–14.4 sec). The mean voluntary dive time of water shrews during 20-min trials ...

Journal of Mammalogy, 2000

Comparative biochemistry and physiology. Part B, Biochemistry & molecular biology, Jan 5, 2018

The high O2 affinity of European mole (Talpa europaea) blood is postulated to largely arise from ... more The high O2 affinity of European mole (Talpa europaea) blood is postulated to largely arise from the presence of two β-globin chain residues (β4 Ser and β5 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 d...

Handbook of experimental pharmacology, Jan 10, 2018

Brown adipose tissue (BAT), the specialized heat-producing organ found in many placental mammals ... more Brown adipose tissue (BAT), the specialized heat-producing organ found in many placental mammals including humans, may be accessible for clinical drug intervention to help combat metabolic diseases. Understanding the biology of BAT and its thermogenic uncoupling protein 1 (UCP1) will benefit from an assessment of its evolution, answering where UCP1 originated and how it has been modified and integrated into cellular energy metabolism. Here, we review topical insights regarding the molecular evolution of UCP1-also reconstructing the proximate and ultimate factors selecting for brown fat thermogenesis in placental mammals. This new thinking on "old" events will assist our understanding of how thermogenic mitochondrial uncoupling was integrated into the physiology of the brown adipocyte. Recent comparative studies examining the occurrence of UCP1 in vertebrates not only identified the ancient (pre-mammal) rise of UCP1 but also its repeated downfall during mammalian evolution ...

Frontiers in physiology, 2018

Zoological research, Jan 28, 2018

Blarinellini is a tribe of soricine shrews comprised of nine fossil genera and one extant genus. ... more Blarinellini is a tribe of soricine shrews comprised of nine fossil genera and one extant genus. Blarinelline shrews were once widely distributed throughout Eurasia and North America, though only members of the Asiatic short-tailed shrew genus Blarinella currently persist (mostly in southwestern China and adjacent areas). Only three forms of Blarinella have been recognized as either species or subspecies. However, recent molecular studies indicated a strikingly deep divergence within the genus, implying the existence of a distinct genus-level lineage. We sequenced the complete mitochondrial genomes and one nuclear gene of three Asiatic short-tailed and two North American shrews and analyzed them morphometrically and morphologically. Our molecular analyses revealed that specimens ascribed to B. griselda formed two deeply diverged lineages, one a close relative to B. quadraticauda, whereas the other - comprised of topotype specimens from southern Gansu - diverged from other Blarinella...

Molecular Phylogenetics and Evolution, 2012

The Journal of Wildlife Management, 1994

... such as forage availability, predation, water level, or suitability for lodge construction ma... more ... such as forage availability, predation, water level, or suitability for lodge construction may have a greater bearing on selection of habitat. ... Like the muskrat, nutria (Myocastor coypus) also feed predominantly on basal shoots and rhi-zomes of aquatic plants, including cattail ...

Journal of Mammalogy, 1996

eLife

The speciose mammalian order Eulipotyphla (moles, shrews, hedgehogs, solenodons) combines an unus... more 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 conf...

eLife

The speciose mammalian order Eulipotyphla (moles, shrews, hedgehogs, solenodons) combines an unus... more 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 conf...

The recently extinct (ca. 1768) Steller's sea cow (Hydrodamalis gigas) was a large, edentulous No... more The recently extinct (ca. 1768) Steller's sea cow (Hydrodamalis gigas) was a large, edentulous North Pacific sirenian. The phylogenetic affinities of this taxon to other members of this clade, living and extinct, are uncertain based on previous morphological and molecular studies. We employed hybridization capture methods and second generation sequencing technology to obtain >30 kb of exon sequences from 26 nuclear genes for both H. gigas and Dugong dugon. We also obtained complete coding sequences for the tooth-related enamelin (ENAM) gene. Hybridization probes designed using dugong and manatee sequences were both highly effective in retrieving sequences from H. gigas (mean = 98.8% coverage), as were more divergent probes for regions of ENAM (99.0% coverage) that were designed exclusively from a proboscidean (African elephant) and a hyracoid (Cape hyrax). New sequences were combined with available sequences for representatives of all other afrotherian orders. We also expanded a previously published morphological matrix for living and fossil Sirenia by adding both new taxa and nine new postcranial characters. Maximum likelihood and parsimony analyses of the molecular data provide robust support for an association of H. gigas and D. dugon to the exclusion of living trichechids (mana-tees). Parsimony analyses of the morphological data also support the inclusion of H. gigas in Dugongidae with D. dugon and fossil dugongids. Timetree analyses based on calibration density approaches with hard-and soft-bounded constraints suggest that H. gigas and D. dugon diverged in the Oligocene and that crown sirenians last shared a common ancestor in the Eocene. The coding sequence for the ENAM gene in H. gigas does not contain frameshift mutations or stop codons, but there is a transversion mutation (AG to CG) in the acceptor splice site of intron 2. This disruption in the edentulous Steller's sea cow is consistent with previous studies that have documented inactivating mutations in tooth-specific loci of a variety of edentulous and enamelless vertebrates including birds, turtles, aardvarks , pangolins, xenarthrans, and baleen whales. Further, branch-site dN/dS analyses provide evidence

The mammalian family Talpidae (moles, shrew moles, desmans) is characterized by diverse ecomorpho... more 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.

Mitochondrial uncoupling protein 1 (UCP1) is essential for nonshivering thermogenesis in brown ad... more Mitochondrial uncoupling protein 1 (UCP1) is essential for nonshivering thermogenesis in brown adipose tissue and is widely accepted to have played a key thermoregulatory role in small-bodied and neonatal placental mammals that enabled the exploitation of cold environments. We map ucp1 sequences from 133 mammals onto a species tree constructed from a ~51-kb sequence alignment and show that inactivating mutations have occurred in at least 8 of the 18 traditional placental orders, thereby challenging the physiological importance of UCP1 across Placentalia. Selection and timetree analyses further reveal that ucp1 inactivations temporally correspond with strong secondary reductions in metabolic intensity in xenarthrans and pangolins, or in six other lineages coincided with a ~30 million–year episode of global cooling in the Paleogene that promoted sharp increases in body mass and cladogenesis evident in the fossil record. Our findings also demonstrate that members of various lineages (for example, cetaceans, horses, woolly mammoths, Steller's sea cows) evolved extreme cold hardiness in the absence of UCP1-mediated thermogenesis. Finally, we identify ucp1 inactivation as a historical contingency that is linked to the current low species diversity of clades lacking functional UCP1, thus providing the first evidence for species selection related to the presence or absence of a single gene product.

Journal of Experimental Biology, 2000

The postprandial increase in metabolic rate associated with consuming, assimilating and excreting... more The postprandial increase in metabolic rate associated with consuming, assimilating and excreting a meal is often termed the heat increment of feeding (HIF). The metabolic heat production of star-nosed moles, Condylura cristata, held at thermoneutrality was monitored for 4 h following a single 10 min session of feeding on a ration consisting of 0 g (controls), 3.5 g or 10 g of earthworms. Coefficients for metabolizable energy digestibility and digesta passage rate of earthworms fed to C. cristata were also determined. We then tested whether feeding-induced thermogenesis substitutes partially or completely for thermoregulatory heat production in these animals exposed to sub-thermoneutral air temperatures (9-24 degrees C). A single feeding on earthworms had both short- and long-term effects on the metabolic rate and respiratory exchange ratio of C. cristata. The observed short-term (0-65 min) rise in metabolic rate, assumed to be associated primarily with the physical costs of nutrien...

ABSTRACTAllometry predicts that the 12–17 g American water shrew (Sorex palustris)—the world’s sm... more ABSTRACTAllometry predicts that the 12–17 g American water shrew (Sorex palustris)—the world’s smallest mammalian diver—will have the highest diving metabolic rate coupled with the lowest total body oxygen storage capacity, skeletal muscle buffering capacity, and glycolytic potential of any endothermic diver. Consistent with expectations, and potentially owing to their low thermal inertia, water shrews had a significantly higher diving metabolic rate in 10°C (8.77 mL O2 g−1 hr−1) compared to 30°C water (6.57 mL O2 g−1 hr−1). Unlike larger-bodied divers, muscle myoglobin contributed minimally (7.7–12.4%) to total onboard O2 stores of juvenile and adult water shrews, respectively, but was offset by high blood O2 carrying capacities (26.4–26.9 vol. %). Diving was predominantly aerobic, as only 2.3–3.9% of dives in 30 and 10°C water, respectively, exceeded the calculated aerobic dive limits at these temperatures (10.8–14.4 sec). The mean voluntary dive time of water shrews during 20-min...

The δ-globin gene (HBD) of eutherian mammals exhibits a propensity for recombinational exchange w... more The δ-globin gene (HBD) of eutherian mammals exhibits a propensity for recombinational exchange with the closely linked β-globin gene (HBB) and has been independently converted by the HBB gene in multiple lineages. Here we report the presence of a chimeric β/δ fusion gene in the African elephant (Loxodonta africana) that was created by unequal crossing-over between misaligned HBD and HBB paralogs. The recombinant chromosome that harbors the β/δ fusion gene in elephants is structurally similar to the ‘‘anti-Lepore’’ duplication mutant of humans (the reciprocal exchange product of the hemoglobin Lepore deletion mutant). However, the situation in the African elephant is unique in that the chimeric β/δ fusion gene supplanted the parental HBB gene and is therefore solely responsible for synthesizing the β-chain subunits of adult hemoglobin. A phylogenetic survey of β-like globin genes in afrotherian and xenarthran mammals revealed that the origin of the chimeric β/δ fusion gene and the c...

bioRxiv, 2021

Allometry predicts that the 12–17 g American water shrew (Sorex palustris)—the world’s smallest m... more Allometry predicts that the 12–17 g American water shrew (Sorex palustris)—the world’s smallest mammalian diver—will have the highest diving metabolic rate coupled with the lowest total body oxygen storage capacity, skeletal muscle buffering capacity, and glycolytic potential of any endothermic diver. Consistent with expectations, and potentially owing to their low thermal inertia, water shrews had a significantly higher diving metabolic rate in 10°C (8.77 mL O2 g−1 hr−1) compared to 30°C water (6.57 mL O2 g−1 hr−1). Unlike larger-bodied divers, muscle myoglobin contributed minimally (7.7–12.4%) to total onboard O2 stores of juvenile and adult water shrews, respectively, but was offset by high blood O2 carrying capacities (26.4–26.9 vol. %). Diving was predominantly aerobic, as only 2.3–3.9% of dives in 30 and 10°C water, respectively, exceeded the calculated aerobic dive limits at these temperatures (10.8–14.4 sec). The mean voluntary dive time of water shrews during 20-min trials ...

Journal of Mammalogy, 2000

Comparative biochemistry and physiology. Part B, Biochemistry & molecular biology, Jan 5, 2018

The high O2 affinity of European mole (Talpa europaea) blood is postulated to largely arise from ... more The high O2 affinity of European mole (Talpa europaea) blood is postulated to largely arise from the presence of two β-globin chain residues (β4 Ser and β5 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 d...

Handbook of experimental pharmacology, Jan 10, 2018

Brown adipose tissue (BAT), the specialized heat-producing organ found in many placental mammals ... more Brown adipose tissue (BAT), the specialized heat-producing organ found in many placental mammals including humans, may be accessible for clinical drug intervention to help combat metabolic diseases. Understanding the biology of BAT and its thermogenic uncoupling protein 1 (UCP1) will benefit from an assessment of its evolution, answering where UCP1 originated and how it has been modified and integrated into cellular energy metabolism. Here, we review topical insights regarding the molecular evolution of UCP1-also reconstructing the proximate and ultimate factors selecting for brown fat thermogenesis in placental mammals. This new thinking on "old" events will assist our understanding of how thermogenic mitochondrial uncoupling was integrated into the physiology of the brown adipocyte. Recent comparative studies examining the occurrence of UCP1 in vertebrates not only identified the ancient (pre-mammal) rise of UCP1 but also its repeated downfall during mammalian evolution ...

Frontiers in physiology, 2018

Zoological research, Jan 28, 2018

Blarinellini is a tribe of soricine shrews comprised of nine fossil genera and one extant genus. ... more Blarinellini is a tribe of soricine shrews comprised of nine fossil genera and one extant genus. Blarinelline shrews were once widely distributed throughout Eurasia and North America, though only members of the Asiatic short-tailed shrew genus Blarinella currently persist (mostly in southwestern China and adjacent areas). Only three forms of Blarinella have been recognized as either species or subspecies. However, recent molecular studies indicated a strikingly deep divergence within the genus, implying the existence of a distinct genus-level lineage. We sequenced the complete mitochondrial genomes and one nuclear gene of three Asiatic short-tailed and two North American shrews and analyzed them morphometrically and morphologically. Our molecular analyses revealed that specimens ascribed to B. griselda formed two deeply diverged lineages, one a close relative to B. quadraticauda, whereas the other - comprised of topotype specimens from southern Gansu - diverged from other Blarinella...

Molecular Phylogenetics and Evolution, 2012

The Journal of Wildlife Management, 1994

... such as forage availability, predation, water level, or suitability for lodge construction ma... more ... such as forage availability, predation, water level, or suitability for lodge construction may have a greater bearing on selection of habitat. ... Like the muskrat, nutria (Myocastor coypus) also feed predominantly on basal shoots and rhi-zomes of aquatic plants, including cattail ...

Journal of Mammalogy, 1996

eLife

The speciose mammalian order Eulipotyphla (moles, shrews, hedgehogs, solenodons) combines an unus... more 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 conf...

eLife

The speciose mammalian order Eulipotyphla (moles, shrews, hedgehogs, solenodons) combines an unus... more 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 conf...

The recently extinct (ca. 1768) Steller's sea cow (Hydrodamalis gigas) was a large, edentulous No... more The recently extinct (ca. 1768) Steller's sea cow (Hydrodamalis gigas) was a large, edentulous North Pacific sirenian. The phylogenetic affinities of this taxon to other members of this clade, living and extinct, are uncertain based on previous morphological and molecular studies. We employed hybridization capture methods and second generation sequencing technology to obtain >30 kb of exon sequences from 26 nuclear genes for both H. gigas and Dugong dugon. We also obtained complete coding sequences for the tooth-related enamelin (ENAM) gene. Hybridization probes designed using dugong and manatee sequences were both highly effective in retrieving sequences from H. gigas (mean = 98.8% coverage), as were more divergent probes for regions of ENAM (99.0% coverage) that were designed exclusively from a proboscidean (African elephant) and a hyracoid (Cape hyrax). New sequences were combined with available sequences for representatives of all other afrotherian orders. We also expanded a previously published morphological matrix for living and fossil Sirenia by adding both new taxa and nine new postcranial characters. Maximum likelihood and parsimony analyses of the molecular data provide robust support for an association of H. gigas and D. dugon to the exclusion of living trichechids (mana-tees). Parsimony analyses of the morphological data also support the inclusion of H. gigas in Dugongidae with D. dugon and fossil dugongids. Timetree analyses based on calibration density approaches with hard-and soft-bounded constraints suggest that H. gigas and D. dugon diverged in the Oligocene and that crown sirenians last shared a common ancestor in the Eocene. The coding sequence for the ENAM gene in H. gigas does not contain frameshift mutations or stop codons, but there is a transversion mutation (AG to CG) in the acceptor splice site of intron 2. This disruption in the edentulous Steller's sea cow is consistent with previous studies that have documented inactivating mutations in tooth-specific loci of a variety of edentulous and enamelless vertebrates including birds, turtles, aardvarks , pangolins, xenarthrans, and baleen whales. Further, branch-site dN/dS analyses provide evidence

The mammalian family Talpidae (moles, shrew moles, desmans) is characterized by diverse ecomorpho... more 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.

Mitochondrial uncoupling protein 1 (UCP1) is essential for nonshivering thermogenesis in brown ad... more Mitochondrial uncoupling protein 1 (UCP1) is essential for nonshivering thermogenesis in brown adipose tissue and is widely accepted to have played a key thermoregulatory role in small-bodied and neonatal placental mammals that enabled the exploitation of cold environments. We map ucp1 sequences from 133 mammals onto a species tree constructed from a ~51-kb sequence alignment and show that inactivating mutations have occurred in at least 8 of the 18 traditional placental orders, thereby challenging the physiological importance of UCP1 across Placentalia. Selection and timetree analyses further reveal that ucp1 inactivations temporally correspond with strong secondary reductions in metabolic intensity in xenarthrans and pangolins, or in six other lineages coincided with a ~30 million–year episode of global cooling in the Paleogene that promoted sharp increases in body mass and cladogenesis evident in the fossil record. Our findings also demonstrate that members of various lineages (for example, cetaceans, horses, woolly mammoths, Steller's sea cows) evolved extreme cold hardiness in the absence of UCP1-mediated thermogenesis. Finally, we identify ucp1 inactivation as a historical contingency that is linked to the current low species diversity of clades lacking functional UCP1, thus providing the first evidence for species selection related to the presence or absence of a single gene product.