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Papers by David Nagorsen

Research paper thumbnail of Mammal collectors' manual

Research paper thumbnail of Port Eliza cave: North American West Coast interstadial environment and implications for human migrations

Quaternary Science Reviews, Jun 1, 2003

The timing of Late Pleistocene glacial advance, retreat, relative sea level and environmental via... more The timing of Late Pleistocene glacial advance, retreat, relative sea level and environmental viability between 25 and 12.5 ka (14 C yrs BP) remain a key issue in the feasibility of a coastal migration route for the first North Americans. This is discussed on the basis of stratigraphic, radiometric and faunal data for Port Eliza cave, a raised sea cave, on the west coast of Vancouver Island, British Columbia. Ice cover is indicated by the occurrence of 2 m of laminated clay, representing deposition in a subglacial lake. From immediately below this clay a diverse vertebrate fauna of marmot, vole, marten, cervid and various species of birds and fish was recovered, yielding ages of 18-16 ka. These dates and others from the region show that ice cover on the outer coast was brief, from ca 15.5-14 ka. The fish species indicate that relative sea level was close to the cave and that salmon runs were likely present. The terrestrial vertebrate fauna is consistent with a cool, open parkland environment with maximum summer temperatures cooler than present; these conditions lasted until at least 16 ka. With a diverse fauna and favorable climate, humans could have survived here on a mixed marine-terrestrial diet, confirming the viability of the coastal migration hypothesis for this portion of the route.

Research paper thumbnail of Phylogeography and Introgressive Hybridization: Chipmunks (Genus Tamias) in the Northern Rocky Mountains

Evolution, 2003

If phylogeographic studies are to be broadly used for assessing population-level processes releva... more If phylogeographic studies are to be broadly used for assessing population-level processes relevant to speciation and systematics, the ability to identify and incorporate instances of hybridization into the analytical framework is essential. Here, we examine the evolutionary history of two chipmunk species, Tamias ruficaudus and Tamias amoenus, in the northern Rocky Mountains by integrating multivariate morphometrics of bacular (os penis) variation, phylogenetic estimation, and nested clade analysis with regional biogeography. Our results indicate multiple examples of mitochondrial DNA introgression layered within the evolutionary history of these nonsister species. Three of these events are most consistent with recent and/or ongoing asymmetric introgression of mitochondrial DNA across morphologically defined secondary contact zones. In addition, we find preliminary evidence where a fourth instance of nonconcordant characters may represent complete fixation of introgressed mitochondrial DNA via a more ancient hybridization event, although alternative explanations of convergence or incomplete sorting of ancestral polymorphisms cannot be dismissed with these data. The demonstration of hybridization among chipmunks with strongly differentiated bacular morphology contradicts long-standing assumptions that variation within this character is diagnostic of complete reproductive isolation within Tamias. Our results illustrate the utility of phylogeographic analyses for detecting instances of reticulate evolution and for incorporating this and other information in the inference of the evolutionary history of species.

Research paper thumbnail of Anoura cultrata

Mammalian Species, Jun 18, 1982

CONTEXT AND CONTENT. Order Chiroptera, Suborder Microchiroptera, Family Phyllostomidae, Subfamily... more CONTEXT AND CONTENT. Order Chiroptera, Suborder Microchiroptera, Family Phyllostomidae, Subfamily Glossophaginae, Genus Anoura. The genus Anoura includes the living species A. caudifer, A. geoffroyi, and A. cultrata (see Diagnosis). A key to living species is as follows: First premolar largest of mandibular premolars, bladelike, strikingly different in shape from other lower premolars in occlusal view Anoura cultrata First lower premolar not enlarged or bladelike, similar to other mandibular premolars in occlusal view 2 2 (1) Interfemoral membrane very narrow, 1.5 to 4 mm wide (about 1.5 to 3 mm in midline), furred dorsally, with a dense fringe of hairs on free margin; forearm length 39 to 48; cranial length 24 to 27; Mexico Eouth to southeastern Brazil and northwestern Argentina, Trinidad, and Grenada A. geoffroyi Interfemoral membrane narrow, 3.5 to 5 mm at midline, wider at level of knees, bearing a few fine hairs forming a thin fringe on free margin; forearm length 34 to 39; cranial length 21 to 25, northern South America south to Peru, Bolivia, and southeastern Brazil ____________________________________________________ A. caudifer Anoura cultrata Handley, 1960 Handley's Long-tongued Bat

Research paper thumbnail of Late Wisconsinan Port Eliza Cave deposits and their implications for human coastal migration, Vancouver Island, Canada

Geoarchaeology, 2006

Sediments of Port Eliza Cave provide a record of the Last Glacial Maximum (LGM) on Vancouver Isla... more Sediments of Port Eliza Cave provide a record of the Last Glacial Maximum (LGM) on Vancouver Island that has important implications for human migration along the debated coastal migration route. Lithofacies changes from nonglacial diamict to glacial laminated silt and clay and till, then a return to nonglacial conditions with oxidized clay, colluvial block beds, and speleothems, along with radiocarbon and U/Th dates, define glacial–nonglacial transitions. Scanning electron microscope studies and clay mineralogy confirm that the laminated fines represent glaciation. Preglacial faunal evidence shows a diverse range from small species, including birds, fish, vole, and marmot, to larger species, such as mountain goat. Pollen data from the same unit show a cold, dry tundra environment with sparse trees. Deglaciation occurred prior to an age of 12.3 ka B.P. based on dated mountain goat bone. These data support the viability of the coastal migration route for humans prior to ˜16 ka B.P. an...

Research paper thumbnail of Mammal collectors' manual

Life Sciences Miscellaneous Publications, Royal Ontario Museum, 1980

The Royal Ontario Museum, 1980 100 Queen's Park, Toronto, Canada M5S 2C6 PRINTED AND BOUND IN CAN... more The Royal Ontario Museum, 1980 100 Queen's Park, Toronto, Canada M5S 2C6 PRINTED AND BOUND IN CANADA AT THE ALGER PRESS measurements, information on weight, sex, age, date, reproductive data, habitat description, and a precise locality description, including latitude and longitude, are stored on magnetic tape. Other institutions with major mammal collections have also begun to use similar computer systems and it is possible that in the future most museum catalogue records may be stored in a central data bank. To utilize fully the potential of these cataloguing systems, collectors should provide the maximum amount of data for specimens. With an increasing concern for the conservation of mammalian species and the additional restrictions being placed on collectors, it is most important that a reasonable and responsible collecting policy be followed. Collecting ethics and collecting laws are discussed in section 2. The recent proliferation of import/export regulations for scientific specimens is another area of concern for collectors. Canadian and US import/export regulations are discussed in some detail in section 7. Collectors are urged to read these two sections carefully before collecting specimens. 2. Collecting Policy 2.1 Collecting Laws In recent years there has been a great increase in the number of collecting laws and endangered species acts that directly affect the scientific collector. These laws may be complex and ambiguous and obtaining the necessary permits for collecting in an area may involve considerable bureaucracy. Nevertheless, the collector has an obligation to learn and comply with these laws and regulations. It is essential that permits be obtained prior to any field collecting. Mammals that are protected under endangered species legislation should not be disturbed or collected except under special permit. Canada In Canada, a Scientific Collector's Licence issued by the various provincial and territorial governments is needed. Fur-bearers and game species are usually regulated under provincial Game Acts and Regulations and special permission may be necessary for collecting these species. Additional permits may be needed to work in a provincial park. In Ontario, for example, collectors who plan to work in provincial parks must have their research proposal approved by the District Manager and the Director of the Parks Branch, Ministry of Natural Resources. Permission from the federal government is required for collecting in national parks. Some provinces and territories also require permits for salvaging dead mammals or parts thereof (carcasses, bones, shed antlers). For more information, consult the appropriate provincial or territorial governments. Although Canada has no federal endangered species act, Canada has signed the Convention on International Trade in Endangered Species (see section 7.2). Ontario and New Brunswick have passed provincial endangered species legislation. United States The US laws are complex and involve state and federal authorities. Generally, wildlife is regulated by the state governments and the collector should contact the appropriate state governmental agency for permits. To collect scientific specimens in a national refuge or in a national park, a permit issued by the Refuge Manager or the Superintendent of the national park is necessary. Permits to collect in national parks are issued only to persons officially representing reputable scientific institutions and annual reports are required. Although state and federal requirements for scientific collecting must be met, special permits are not needed for collecting in national forest systems. Collectors, however, are requested to contact the local Forest Service District Ranger before initiating any fieldwork. Marine mammals are covered under the Marine Mammal Protection Act. You may take and process marine mammals and parts thereof (bones, teeth, ivory) only under permit. Walruses (Qdobenus rosmarus), sea otters (Enhydra lutris), polar bears (Ursus maritimus), and manatees (Trichechus sp.) are under the jurisdiction of the Director of the US Fish and Wildlife Service. Cetaceans and all pinnipeds (except walruses) are under the authority of the Director of the National Marine Fisheries Service. Collectors working in the US should be aware that endangered and threatened species are protected at both the federal and state levels. The federal Endangered Species Act that took effect in 1977 prohibits the taking and capture of all listed mammals as well as the import and export of these species. Species protected under the Act are listed as either "endangered" or "threatened". The various prohibitions of the Act apply to live or dead mammals and their parts or products. If you plan to salvage or utilize dead mammals listed in the Act, you must have a permit issued by the Director of the US Fish and Wildlife Service, or if endangered marine mammals, from the Director of National Marine Fisheries Service. For further information, contact the Federal Wildlife Permit Office, US Fish and Wildlife Service, Washington, DC 20240. Some states have also passed endangered species legislation and mammals protected under state law may or may not be the same as those listed under the federal Endangered Species Act. Information on these state laws can be obtained from the appropriate state governmental authorities such as conservation or wildlife departments (see McGaugh and Genoways 1976). Other Countries Many other countries have also passed endangered species legislation and regulations for the collecting of scientific specimens. Collectors planning to work abroad should contact the governmental agencies in these countries well in advance of any field trip. It is essential that one understand the regulations in these countries and obtain the necessary permits before any field collecting. Moreover, export permits may be required in some countries for transporting specimens out of these countries. The collector should also be aware of the Convention on International Trade in Endangered Species. Even if species listed in the Convention are not protected in the country where the collector is working, it may be impossible to import specimens of species listed in the Convention into North America without permits (see section 7). 2.2 Firearms In recent years there has been a growing anti-hunting sentiment and collectors therefore are urged to use discretion when collecting with guns. Handguns are strictly controlled in Canada and permits issued by police departments are required. In some provinces municipal or provincial police departments may issue permits; in other provinces, the Yukon, and the Northwest Territories, permits are issued by the Royal Canadian Mounted Police (RCMP). In the US, firearms are generally regulated by the various state governments and collectors should consult their state or local police departments for information. When planning to use firearms in foreign countries, investigate thoroughly the various laws in these countries pertaining to firearms and ammunition. These laws include customs regulations covering the importation of guns and ammunition. 2.3 Collecting Ethics The possession of a valid collecting permit does not give the collector the right to use irresponsible collecting methods. Specimens should be collected in the most humane method possible and any damage or destruction to the local biota or collecting sites must be prevented. Indiscriminate collecting of excessive numbers is discouraged, particularly in areas where large numbers may concentrate. When possible, take mammals alive and once the required sample has been collected, release the remaining mammals unharmed. Obviously with techniques such as snap trapping this procedure is impossible to follow. Specimens acquired for systematic collections should be carefully prepared and thoroughly documented using the standards outlined in this manual. Systematic research on a species requires a statistically adequate number of specimens from various localities. Because some species are sexually dimorphic (e.g., one sex may be consistently larger than the other), a representative series should contain samples of both sexes. For a given locality, 10 to 15 adults of each sex are usually an adequate number for a species. A few young animals in each sample may be useful for studying growth and age variation. To study geographic variation in a species, representative samples from various localities throughout the geographic range of the species are required. Distance between collecting sites is a function of habitat diversity in a given area and in regions with homogeneous biomes and habitats (e.g., the boreal forest in northern Canada or the tropical rain forest in the Amazon basin) localities may be 15 to 160 km (10-100 miles) apart. But in regions that support a diversity of biomes and habitats (e.g., mountainous regions or river systems), collecting sites may need to be close together, 8 km (5 miles) or less. 3. Methods for Collecting Mammals 3.1 Bats A. Equipment mist NETS Mist nets or bird-banders' nets (Bleitz Wildlife Foundation, 5334 Hollywood Boulevard, Hollywood, California, USA) are effective for capturing bats alive. The

Research paper thumbnail of Conserving Mammals at Risk : The Role of Taxonomy

A number of mammalian species, subspecies, and populations are designated at risk both nationally... more A number of mammalian species, subspecies, and populations are designated at risk both nationally and by the province of British Columbia. Despite the perception that the taxonomy of mammals is well known, major taxonomic problems are associated with some of these taxa. Species limits and reliable identification traits need to be determined for several problematic species groups. Moreover, the validity of many of the subspecies listed at risk is unknown. Some were described more than 50 years ago from small samples and descriptive methodology, and modern research is essential to determine if they represent distinct evolutionary units that warrant protection. We use recent research on listed subspecies of chipmunks (Tamias spp.), northern pocket gophers (Thomomys talpoides), and southern red-backed voles (Clethrionomys gapperi) from the Kootenay region of British Columbia to illustrate how taxonomy is critical for delimiting conservation units. Preliminary results suggest that the 2 ...

Research paper thumbnail of Reply to the comment by Morales et al. on “Population genetics reveal Myotis keenii (Keen’s myotis) and Myotis evotis (long-eared myotis) to be a single species”

Canadian Journal of Zoology, 2021

A.E. Morales et al. (2021. Can. J. Zool. 99(5): 415–422) provided no new evidence to alter the co... more A.E. Morales et al. (2021. Can. J. Zool. 99(5): 415–422) provided no new evidence to alter the conclusions of C.L. Lausen et al. (2019. Can. J. Zool. 97(3): 267–279). We present background information, relevant comparisons, and clarification of analyses to further strengthen our conclusions. The genesis of the original “evotis–keenii” study in British Columbia (Canada) was to differentiate Myotis keenii (Merriam, 1895) (Keen’s myotis), with one of the smallest North American bat distributions, from sympatric Myotis evotis (H. Allen, 1864) (long-eared myotis), using something other than the suggested post-mortem skull size comparison, but no differentiating trait could be found, leading to the molecular genetics examination of C.L. Lausen et al. (2019). We present cumulative data that rejects the 1979 hypothesis of M. keenii as a distinct species. A.E. Morales et al. (2021) inaccurately portray C.L. Lausen et al.’s (2019) question and results; present inaccurate morphological and out...

Research paper thumbnail of Karyotypes of six species of bats (Chiroptera) from the Dominican Republic /

Karyotypes of Mormoops blainvilli, Macrotus w. waterhousii, Phyllops haitiensis, Brachyphylla pum... more Karyotypes of Mormoops blainvilli, Macrotus w. waterhousii, Phyllops haitiensis, Brachyphylla pumila, Phyllonycteris phology were determined from photographs of six or more metaphase spreads of each species. Voucher specimens are in the collections of the Department of Mammalogy, Royal Ontario Museum (rom). department of Mammalogy, Royal Ontario Museum.

Research paper thumbnail of Systematics of Anoura cultrata, A. brevirostrum, and A. werckleae

Journal of Mammalogy, 1981

... DAVID NAGORSEN AND JR TAMSITT ... from maps of plant formations for Peru (Tosi, 1957), Colomb... more ... DAVID NAGORSEN AND JR TAMSITT ... from maps of plant formations for Peru (Tosi, 1957), Colombia (Espinal and Montenegro, 1963), Venezuela (Ewel and Madriz, 1968; Handley, 1976), Panama (Holdridge and Budowski, 1959), and Costa Rica (Holdridge, 1959). ... A .0 c -.2-...

Research paper thumbnail of Phenotypes and Distribution of Yellow-Pine Chipmunk (Neotamias amoenus) of Hybrid Ancestry from the Rocky Mountains of Canada

Western North American Naturalist, 2021

Abstract. In Canada, subspecies of the yellow-pine chipmunk (Neotamias amoenus luteiventris) and ... more Abstract. In Canada, subspecies of the yellow-pine chipmunk (Neotamias amoenus luteiventris) and red-tailed chipmunk (Neotamias ruficaudus ruficaudus) co-occur in a small area of the Rocky Mountains in southwestern Alberta and southeastern British Columbia. Some N. a. luteiventris individuals in this area are of hybrid ancestry, carrying a mitochondrial DNA gene of N. r. ruficaudus. We tested for phenotypic hybridization by comparing genital bone morphology, body size, cranial mandibular morphology, and tail color of mitochondrial hybrids with specimens of the 2 parental reference groups identified from genetics or genital morphology. Hybrids were indistinguishable from the N. a. luteiventris reference group, demonstrating no intermediacy or increased variability. Our results are consistent with genetic data that suggest minimal nuclear gene flow between the 2 taxa. Five hybrid locations were within the contact zone, but 6 were 14–98 km beyond the known range extent of N. r. ruficaudus. These peripheral occurrences may be a legacy of a past range decline of N. r. ruficaudus in response to Holocene climatic changes that left hybrids outside the present-day contact zone. We recommend more field research in the Canadian Rocky Mountains to determine the full range extent of N. r. ruficaudus and N. a. luteiventris of hybrid ancestry.

Research paper thumbnail of Population genetics reveal Myotis keenii (Keen’s myotis) and Myotis evotis (long-eared myotis) to be a single species

Canadian Journal of Zoology

Recognizing delineations of gene flow among groups of animals can be challenging but is necessary... more Recognizing delineations of gene flow among groups of animals can be challenging but is necessary for conservation and management. Of particular importance is the identification of species boundaries. Several physical and genetic traits have been used with mixed success to distinguish Myotis keenii (Merriam, 1895) (Keen’s myotis) and Myotis evotis (H. Allen, 1864) (long-eared myotis), but it is unclear whether species distinction is biologically warranted. We generated 12–14 microsatellite locus genotypes for 275 long-eared Myotis representing four species — M. keenii, M. evotis, Myotis septentrionalis (Trouessart, 1897) (northern myotis), and Myotis thysanodes Miller, 1897 (fringed myotis) — from across northwestern North America and 23 Myotis lucifugus (Le Conte, 1831) (little brown myotis) as the outgroup. Population genetic analyses revealed four well-defined groups (species): M. septentrionalis, M. thysanodes, M. lucifugus, and a single group comprising M. keenii and M. evotis....

Research paper thumbnail of Melanistic Tundra Voles, <em>Microtus oeconomus</em>, from Central Yukon

The Canadian Field-Naturalist

Colour aberrations are not commonly observed in voles (e.g., Microtus and Myodes); thus, individu... more Colour aberrations are not commonly observed in voles (e.g., Microtus and Myodes); thus, individual observations are of interest. We report two observations of melanism in Tundra Voles, Microtus oeconomus, collected from central Yukon. These are the second and third records of melanistic Tundra Voles, and the first reports from non-insular populations.

Research paper thumbnail of Northern Range Extension of the Pygmy Shrew, <em>Sorex hoyi</em>, in the Yukon

The Canadian Field-Naturalist

A Pygmy Shrew, Sorex hoyi, was captured in a pitfall trap on the Blackstone River (65°04.6'N,... more A Pygmy Shrew, Sorex hoyi, was captured in a pitfall trap on the Blackstone River (65°04.6'N, 138°10.8'W) in central Yukon. This represents a northern range extension of about 110 km for S. hoyi in the Yukon.

Research paper thumbnail of First Records of the Northern Long-eared Bat, <em>Myotis septentrionalis</em>, in the Yukon Territory

The Canadian Field-Naturalist

Three adult male Northern Long-eared Bats, Myotis septentrionalis, were captured in mist nets in ... more Three adult male Northern Long-eared Bats, Myotis septentrionalis, were captured in mist nets in July 2004 in the LaBiche River Valley, southeastern Yukon. These are the first records of M. septentrionalis in the Yukon. Further survey work is needed to delineate the extent of the range and population structure of this and other species of bats in northwestern North America.

Research paper thumbnail of Are the western water shrew (Sorex navigator) and American water shrew (Sorex palustris) morphologically distinct?

Canadian Journal of Zoology

Recent phylogenetic studies applying the mitochondrial cytochrome b (Cytb) gene and nuclear genes... more Recent phylogenetic studies applying the mitochondrial cytochrome b (Cytb) gene and nuclear genes revealed divergent cordilleran and boreal lineages in western populations of the Sorex palustris complex that are now recognized as distinct species: Sorex navigator (Baird, 1858) (western water shrew) and Sorex palustris Richardson, 1828 (American water shrew). Their distributions in western Canada are parapatric with a potential contact zone in the foothills of the Rocky Mountains. Using 80 specimens including 69 of known genotype, we assessed morphological differentiation in metric and nonmetric traits among S. navigator, S. palustris, and Sorex bendirii (Merriam, 1894) (marsh shrew), a sister species of S. navigator. Multivariate analyses revealed discrete morpho groups concordant with their genetic lineages and species-level divergence. Three individuals from northwestern British Columbia and one from southwestern Alberta with discordant mitochondrial and nuclear DNA aligned with t...

Research paper thumbnail of New records of the Ogilvie Mountains Collared Lemming (<em>Dicrostonyx nunatakensis</em>) in central Yukon

The Canadian Field-Naturalist, 2014

The Ogilvie Mountain Collared Lemming (Dicrostonyx nunatakensis Youngman, 1967), reported only fr... more The Ogilvie Mountain Collared Lemming (Dicrostonyx nunatakensis Youngman, 1967), reported only from the Ogilvie Mountains of central Yukon, is among the least known mammals in Canada. It was first discovered in 1961 and, since then, only 13 specimens had been collected, all from one mountain, in central Yukon. We conducted a targeted survey to determine the distribution of the species by trapping areas of apparently suitable habitat on 12 mountains within 40 km of the known location. Many of our traps were disabled by other mammals; however, we captured three Ogilvie Mountain Collared Lemmings on two mountains 25.9 km and 29.6 km from the original location. Our findings suggest that this lemming may be more widely distributed than indicated by earlier specimens. We suggest further surveys to delineate the range of the Ogilvie Mountain Collared Lemming.

Research paper thumbnail of A Tribute to Ian McTaggart-Cowan, 1910-2010, O.C., O.B.C., PhD, LL.D, F.R.S.C

The Canadian Field-Naturalist, 2010

Research paper thumbnail of First Records of the Southern Red-backed Vole, <em>Myodes gapperi</em>, in the Yukon

The Canadian Field-Naturalist, 2006

Twenty Southern Red-backed Voles, Myodes gapperi, were collected in July 2004 in the LaBiche Rive... more Twenty Southern Red-backed Voles, Myodes gapperi, were collected in July 2004 in the LaBiche River valley of southeastern Yukon. Specimens were identified using morphological characteristics and analysis of cytochrome b gene sequences. These are the first records of this species in the Yukon. No Northern Red-backed Voles, M. rutilus, were collected and it is not known whether the two species are sympatric or parapatric in the Yukon. Further survey work is needed in southeastern Yukon to better delineate the extent of the northwestern range of this species and the extent, if any, of introgression with M. rutilus.

Research paper thumbnail of A morphometric study of geographic variation in the snowshoe hare (Lepus americanus)

Canadian Journal of Zoology, Feb 14, 2011

Geographic variation in Lepus americanus was studied by multivariate analyses of 15 cranial measu... more Geographic variation in Lepus americanus was studied by multivariate analyses of 15 cranial measurements. A total of 1494 specimens from the entire geographic range were grouped into 37 geographic samples of males and females. Principal component analyses demonstrated that hares are largest in eastern North America, Alaska, and northwestern Canada, and smallest in the Pacific Northwest. Size clines exist in the Appalachian Mountains and western North America but size is relatively uniform throughout central Canada and the Great Lakes. A multiple regression of size with 16 climatic variables factors demonstrated that size and climate are strongly correlated; the size trends may reflect environmental selection. The differentiation of hares from the western Cordillera and Pacific coast, the similarity of populations from central Canada and the Great Lakes, and the clines in the Appalachians and western North America were evident in discriminant analyses. These patterns of variation among populations can be attributed to both gene flow and local selection. There is no evidence from the morphometric analyses for classifying populations into the 15 subspecies currently recognized.

Research paper thumbnail of Mammal collectors' manual

Research paper thumbnail of Port Eliza cave: North American West Coast interstadial environment and implications for human migrations

Quaternary Science Reviews, Jun 1, 2003

The timing of Late Pleistocene glacial advance, retreat, relative sea level and environmental via... more The timing of Late Pleistocene glacial advance, retreat, relative sea level and environmental viability between 25 and 12.5 ka (14 C yrs BP) remain a key issue in the feasibility of a coastal migration route for the first North Americans. This is discussed on the basis of stratigraphic, radiometric and faunal data for Port Eliza cave, a raised sea cave, on the west coast of Vancouver Island, British Columbia. Ice cover is indicated by the occurrence of 2 m of laminated clay, representing deposition in a subglacial lake. From immediately below this clay a diverse vertebrate fauna of marmot, vole, marten, cervid and various species of birds and fish was recovered, yielding ages of 18-16 ka. These dates and others from the region show that ice cover on the outer coast was brief, from ca 15.5-14 ka. The fish species indicate that relative sea level was close to the cave and that salmon runs were likely present. The terrestrial vertebrate fauna is consistent with a cool, open parkland environment with maximum summer temperatures cooler than present; these conditions lasted until at least 16 ka. With a diverse fauna and favorable climate, humans could have survived here on a mixed marine-terrestrial diet, confirming the viability of the coastal migration hypothesis for this portion of the route.

Research paper thumbnail of Phylogeography and Introgressive Hybridization: Chipmunks (Genus Tamias) in the Northern Rocky Mountains

Evolution, 2003

If phylogeographic studies are to be broadly used for assessing population-level processes releva... more If phylogeographic studies are to be broadly used for assessing population-level processes relevant to speciation and systematics, the ability to identify and incorporate instances of hybridization into the analytical framework is essential. Here, we examine the evolutionary history of two chipmunk species, Tamias ruficaudus and Tamias amoenus, in the northern Rocky Mountains by integrating multivariate morphometrics of bacular (os penis) variation, phylogenetic estimation, and nested clade analysis with regional biogeography. Our results indicate multiple examples of mitochondrial DNA introgression layered within the evolutionary history of these nonsister species. Three of these events are most consistent with recent and/or ongoing asymmetric introgression of mitochondrial DNA across morphologically defined secondary contact zones. In addition, we find preliminary evidence where a fourth instance of nonconcordant characters may represent complete fixation of introgressed mitochondrial DNA via a more ancient hybridization event, although alternative explanations of convergence or incomplete sorting of ancestral polymorphisms cannot be dismissed with these data. The demonstration of hybridization among chipmunks with strongly differentiated bacular morphology contradicts long-standing assumptions that variation within this character is diagnostic of complete reproductive isolation within Tamias. Our results illustrate the utility of phylogeographic analyses for detecting instances of reticulate evolution and for incorporating this and other information in the inference of the evolutionary history of species.

Research paper thumbnail of Anoura cultrata

Mammalian Species, Jun 18, 1982

CONTEXT AND CONTENT. Order Chiroptera, Suborder Microchiroptera, Family Phyllostomidae, Subfamily... more CONTEXT AND CONTENT. Order Chiroptera, Suborder Microchiroptera, Family Phyllostomidae, Subfamily Glossophaginae, Genus Anoura. The genus Anoura includes the living species A. caudifer, A. geoffroyi, and A. cultrata (see Diagnosis). A key to living species is as follows: First premolar largest of mandibular premolars, bladelike, strikingly different in shape from other lower premolars in occlusal view Anoura cultrata First lower premolar not enlarged or bladelike, similar to other mandibular premolars in occlusal view 2 2 (1) Interfemoral membrane very narrow, 1.5 to 4 mm wide (about 1.5 to 3 mm in midline), furred dorsally, with a dense fringe of hairs on free margin; forearm length 39 to 48; cranial length 24 to 27; Mexico Eouth to southeastern Brazil and northwestern Argentina, Trinidad, and Grenada A. geoffroyi Interfemoral membrane narrow, 3.5 to 5 mm at midline, wider at level of knees, bearing a few fine hairs forming a thin fringe on free margin; forearm length 34 to 39; cranial length 21 to 25, northern South America south to Peru, Bolivia, and southeastern Brazil ____________________________________________________ A. caudifer Anoura cultrata Handley, 1960 Handley's Long-tongued Bat

Research paper thumbnail of Late Wisconsinan Port Eliza Cave deposits and their implications for human coastal migration, Vancouver Island, Canada

Geoarchaeology, 2006

Sediments of Port Eliza Cave provide a record of the Last Glacial Maximum (LGM) on Vancouver Isla... more Sediments of Port Eliza Cave provide a record of the Last Glacial Maximum (LGM) on Vancouver Island that has important implications for human migration along the debated coastal migration route. Lithofacies changes from nonglacial diamict to glacial laminated silt and clay and till, then a return to nonglacial conditions with oxidized clay, colluvial block beds, and speleothems, along with radiocarbon and U/Th dates, define glacial–nonglacial transitions. Scanning electron microscope studies and clay mineralogy confirm that the laminated fines represent glaciation. Preglacial faunal evidence shows a diverse range from small species, including birds, fish, vole, and marmot, to larger species, such as mountain goat. Pollen data from the same unit show a cold, dry tundra environment with sparse trees. Deglaciation occurred prior to an age of 12.3 ka B.P. based on dated mountain goat bone. These data support the viability of the coastal migration route for humans prior to ˜16 ka B.P. an...

Research paper thumbnail of Mammal collectors' manual

Life Sciences Miscellaneous Publications, Royal Ontario Museum, 1980

The Royal Ontario Museum, 1980 100 Queen's Park, Toronto, Canada M5S 2C6 PRINTED AND BOUND IN CAN... more The Royal Ontario Museum, 1980 100 Queen's Park, Toronto, Canada M5S 2C6 PRINTED AND BOUND IN CANADA AT THE ALGER PRESS measurements, information on weight, sex, age, date, reproductive data, habitat description, and a precise locality description, including latitude and longitude, are stored on magnetic tape. Other institutions with major mammal collections have also begun to use similar computer systems and it is possible that in the future most museum catalogue records may be stored in a central data bank. To utilize fully the potential of these cataloguing systems, collectors should provide the maximum amount of data for specimens. With an increasing concern for the conservation of mammalian species and the additional restrictions being placed on collectors, it is most important that a reasonable and responsible collecting policy be followed. Collecting ethics and collecting laws are discussed in section 2. The recent proliferation of import/export regulations for scientific specimens is another area of concern for collectors. Canadian and US import/export regulations are discussed in some detail in section 7. Collectors are urged to read these two sections carefully before collecting specimens. 2. Collecting Policy 2.1 Collecting Laws In recent years there has been a great increase in the number of collecting laws and endangered species acts that directly affect the scientific collector. These laws may be complex and ambiguous and obtaining the necessary permits for collecting in an area may involve considerable bureaucracy. Nevertheless, the collector has an obligation to learn and comply with these laws and regulations. It is essential that permits be obtained prior to any field collecting. Mammals that are protected under endangered species legislation should not be disturbed or collected except under special permit. Canada In Canada, a Scientific Collector's Licence issued by the various provincial and territorial governments is needed. Fur-bearers and game species are usually regulated under provincial Game Acts and Regulations and special permission may be necessary for collecting these species. Additional permits may be needed to work in a provincial park. In Ontario, for example, collectors who plan to work in provincial parks must have their research proposal approved by the District Manager and the Director of the Parks Branch, Ministry of Natural Resources. Permission from the federal government is required for collecting in national parks. Some provinces and territories also require permits for salvaging dead mammals or parts thereof (carcasses, bones, shed antlers). For more information, consult the appropriate provincial or territorial governments. Although Canada has no federal endangered species act, Canada has signed the Convention on International Trade in Endangered Species (see section 7.2). Ontario and New Brunswick have passed provincial endangered species legislation. United States The US laws are complex and involve state and federal authorities. Generally, wildlife is regulated by the state governments and the collector should contact the appropriate state governmental agency for permits. To collect scientific specimens in a national refuge or in a national park, a permit issued by the Refuge Manager or the Superintendent of the national park is necessary. Permits to collect in national parks are issued only to persons officially representing reputable scientific institutions and annual reports are required. Although state and federal requirements for scientific collecting must be met, special permits are not needed for collecting in national forest systems. Collectors, however, are requested to contact the local Forest Service District Ranger before initiating any fieldwork. Marine mammals are covered under the Marine Mammal Protection Act. You may take and process marine mammals and parts thereof (bones, teeth, ivory) only under permit. Walruses (Qdobenus rosmarus), sea otters (Enhydra lutris), polar bears (Ursus maritimus), and manatees (Trichechus sp.) are under the jurisdiction of the Director of the US Fish and Wildlife Service. Cetaceans and all pinnipeds (except walruses) are under the authority of the Director of the National Marine Fisheries Service. Collectors working in the US should be aware that endangered and threatened species are protected at both the federal and state levels. The federal Endangered Species Act that took effect in 1977 prohibits the taking and capture of all listed mammals as well as the import and export of these species. Species protected under the Act are listed as either "endangered" or "threatened". The various prohibitions of the Act apply to live or dead mammals and their parts or products. If you plan to salvage or utilize dead mammals listed in the Act, you must have a permit issued by the Director of the US Fish and Wildlife Service, or if endangered marine mammals, from the Director of National Marine Fisheries Service. For further information, contact the Federal Wildlife Permit Office, US Fish and Wildlife Service, Washington, DC 20240. Some states have also passed endangered species legislation and mammals protected under state law may or may not be the same as those listed under the federal Endangered Species Act. Information on these state laws can be obtained from the appropriate state governmental authorities such as conservation or wildlife departments (see McGaugh and Genoways 1976). Other Countries Many other countries have also passed endangered species legislation and regulations for the collecting of scientific specimens. Collectors planning to work abroad should contact the governmental agencies in these countries well in advance of any field trip. It is essential that one understand the regulations in these countries and obtain the necessary permits before any field collecting. Moreover, export permits may be required in some countries for transporting specimens out of these countries. The collector should also be aware of the Convention on International Trade in Endangered Species. Even if species listed in the Convention are not protected in the country where the collector is working, it may be impossible to import specimens of species listed in the Convention into North America without permits (see section 7). 2.2 Firearms In recent years there has been a growing anti-hunting sentiment and collectors therefore are urged to use discretion when collecting with guns. Handguns are strictly controlled in Canada and permits issued by police departments are required. In some provinces municipal or provincial police departments may issue permits; in other provinces, the Yukon, and the Northwest Territories, permits are issued by the Royal Canadian Mounted Police (RCMP). In the US, firearms are generally regulated by the various state governments and collectors should consult their state or local police departments for information. When planning to use firearms in foreign countries, investigate thoroughly the various laws in these countries pertaining to firearms and ammunition. These laws include customs regulations covering the importation of guns and ammunition. 2.3 Collecting Ethics The possession of a valid collecting permit does not give the collector the right to use irresponsible collecting methods. Specimens should be collected in the most humane method possible and any damage or destruction to the local biota or collecting sites must be prevented. Indiscriminate collecting of excessive numbers is discouraged, particularly in areas where large numbers may concentrate. When possible, take mammals alive and once the required sample has been collected, release the remaining mammals unharmed. Obviously with techniques such as snap trapping this procedure is impossible to follow. Specimens acquired for systematic collections should be carefully prepared and thoroughly documented using the standards outlined in this manual. Systematic research on a species requires a statistically adequate number of specimens from various localities. Because some species are sexually dimorphic (e.g., one sex may be consistently larger than the other), a representative series should contain samples of both sexes. For a given locality, 10 to 15 adults of each sex are usually an adequate number for a species. A few young animals in each sample may be useful for studying growth and age variation. To study geographic variation in a species, representative samples from various localities throughout the geographic range of the species are required. Distance between collecting sites is a function of habitat diversity in a given area and in regions with homogeneous biomes and habitats (e.g., the boreal forest in northern Canada or the tropical rain forest in the Amazon basin) localities may be 15 to 160 km (10-100 miles) apart. But in regions that support a diversity of biomes and habitats (e.g., mountainous regions or river systems), collecting sites may need to be close together, 8 km (5 miles) or less. 3. Methods for Collecting Mammals 3.1 Bats A. Equipment mist NETS Mist nets or bird-banders' nets (Bleitz Wildlife Foundation, 5334 Hollywood Boulevard, Hollywood, California, USA) are effective for capturing bats alive. The

Research paper thumbnail of Conserving Mammals at Risk : The Role of Taxonomy

A number of mammalian species, subspecies, and populations are designated at risk both nationally... more A number of mammalian species, subspecies, and populations are designated at risk both nationally and by the province of British Columbia. Despite the perception that the taxonomy of mammals is well known, major taxonomic problems are associated with some of these taxa. Species limits and reliable identification traits need to be determined for several problematic species groups. Moreover, the validity of many of the subspecies listed at risk is unknown. Some were described more than 50 years ago from small samples and descriptive methodology, and modern research is essential to determine if they represent distinct evolutionary units that warrant protection. We use recent research on listed subspecies of chipmunks (Tamias spp.), northern pocket gophers (Thomomys talpoides), and southern red-backed voles (Clethrionomys gapperi) from the Kootenay region of British Columbia to illustrate how taxonomy is critical for delimiting conservation units. Preliminary results suggest that the 2 ...

Research paper thumbnail of Reply to the comment by Morales et al. on “Population genetics reveal Myotis keenii (Keen’s myotis) and Myotis evotis (long-eared myotis) to be a single species”

Canadian Journal of Zoology, 2021

A.E. Morales et al. (2021. Can. J. Zool. 99(5): 415–422) provided no new evidence to alter the co... more A.E. Morales et al. (2021. Can. J. Zool. 99(5): 415–422) provided no new evidence to alter the conclusions of C.L. Lausen et al. (2019. Can. J. Zool. 97(3): 267–279). We present background information, relevant comparisons, and clarification of analyses to further strengthen our conclusions. The genesis of the original “evotis–keenii” study in British Columbia (Canada) was to differentiate Myotis keenii (Merriam, 1895) (Keen’s myotis), with one of the smallest North American bat distributions, from sympatric Myotis evotis (H. Allen, 1864) (long-eared myotis), using something other than the suggested post-mortem skull size comparison, but no differentiating trait could be found, leading to the molecular genetics examination of C.L. Lausen et al. (2019). We present cumulative data that rejects the 1979 hypothesis of M. keenii as a distinct species. A.E. Morales et al. (2021) inaccurately portray C.L. Lausen et al.’s (2019) question and results; present inaccurate morphological and out...

Research paper thumbnail of Karyotypes of six species of bats (Chiroptera) from the Dominican Republic /

Karyotypes of Mormoops blainvilli, Macrotus w. waterhousii, Phyllops haitiensis, Brachyphylla pum... more Karyotypes of Mormoops blainvilli, Macrotus w. waterhousii, Phyllops haitiensis, Brachyphylla pumila, Phyllonycteris phology were determined from photographs of six or more metaphase spreads of each species. Voucher specimens are in the collections of the Department of Mammalogy, Royal Ontario Museum (rom). department of Mammalogy, Royal Ontario Museum.

Research paper thumbnail of Systematics of Anoura cultrata, A. brevirostrum, and A. werckleae

Journal of Mammalogy, 1981

... DAVID NAGORSEN AND JR TAMSITT ... from maps of plant formations for Peru (Tosi, 1957), Colomb... more ... DAVID NAGORSEN AND JR TAMSITT ... from maps of plant formations for Peru (Tosi, 1957), Colombia (Espinal and Montenegro, 1963), Venezuela (Ewel and Madriz, 1968; Handley, 1976), Panama (Holdridge and Budowski, 1959), and Costa Rica (Holdridge, 1959). ... A .0 c -.2-...

Research paper thumbnail of Phenotypes and Distribution of Yellow-Pine Chipmunk (Neotamias amoenus) of Hybrid Ancestry from the Rocky Mountains of Canada

Western North American Naturalist, 2021

Abstract. In Canada, subspecies of the yellow-pine chipmunk (Neotamias amoenus luteiventris) and ... more Abstract. In Canada, subspecies of the yellow-pine chipmunk (Neotamias amoenus luteiventris) and red-tailed chipmunk (Neotamias ruficaudus ruficaudus) co-occur in a small area of the Rocky Mountains in southwestern Alberta and southeastern British Columbia. Some N. a. luteiventris individuals in this area are of hybrid ancestry, carrying a mitochondrial DNA gene of N. r. ruficaudus. We tested for phenotypic hybridization by comparing genital bone morphology, body size, cranial mandibular morphology, and tail color of mitochondrial hybrids with specimens of the 2 parental reference groups identified from genetics or genital morphology. Hybrids were indistinguishable from the N. a. luteiventris reference group, demonstrating no intermediacy or increased variability. Our results are consistent with genetic data that suggest minimal nuclear gene flow between the 2 taxa. Five hybrid locations were within the contact zone, but 6 were 14–98 km beyond the known range extent of N. r. ruficaudus. These peripheral occurrences may be a legacy of a past range decline of N. r. ruficaudus in response to Holocene climatic changes that left hybrids outside the present-day contact zone. We recommend more field research in the Canadian Rocky Mountains to determine the full range extent of N. r. ruficaudus and N. a. luteiventris of hybrid ancestry.

Research paper thumbnail of Population genetics reveal Myotis keenii (Keen’s myotis) and Myotis evotis (long-eared myotis) to be a single species

Canadian Journal of Zoology

Recognizing delineations of gene flow among groups of animals can be challenging but is necessary... more Recognizing delineations of gene flow among groups of animals can be challenging but is necessary for conservation and management. Of particular importance is the identification of species boundaries. Several physical and genetic traits have been used with mixed success to distinguish Myotis keenii (Merriam, 1895) (Keen’s myotis) and Myotis evotis (H. Allen, 1864) (long-eared myotis), but it is unclear whether species distinction is biologically warranted. We generated 12–14 microsatellite locus genotypes for 275 long-eared Myotis representing four species — M. keenii, M. evotis, Myotis septentrionalis (Trouessart, 1897) (northern myotis), and Myotis thysanodes Miller, 1897 (fringed myotis) — from across northwestern North America and 23 Myotis lucifugus (Le Conte, 1831) (little brown myotis) as the outgroup. Population genetic analyses revealed four well-defined groups (species): M. septentrionalis, M. thysanodes, M. lucifugus, and a single group comprising M. keenii and M. evotis....

Research paper thumbnail of Melanistic Tundra Voles, <em>Microtus oeconomus</em>, from Central Yukon

The Canadian Field-Naturalist

Colour aberrations are not commonly observed in voles (e.g., Microtus and Myodes); thus, individu... more Colour aberrations are not commonly observed in voles (e.g., Microtus and Myodes); thus, individual observations are of interest. We report two observations of melanism in Tundra Voles, Microtus oeconomus, collected from central Yukon. These are the second and third records of melanistic Tundra Voles, and the first reports from non-insular populations.

Research paper thumbnail of Northern Range Extension of the Pygmy Shrew, <em>Sorex hoyi</em>, in the Yukon

The Canadian Field-Naturalist

A Pygmy Shrew, Sorex hoyi, was captured in a pitfall trap on the Blackstone River (65°04.6'N,... more A Pygmy Shrew, Sorex hoyi, was captured in a pitfall trap on the Blackstone River (65°04.6'N, 138°10.8'W) in central Yukon. This represents a northern range extension of about 110 km for S. hoyi in the Yukon.

Research paper thumbnail of First Records of the Northern Long-eared Bat, <em>Myotis septentrionalis</em>, in the Yukon Territory

The Canadian Field-Naturalist

Three adult male Northern Long-eared Bats, Myotis septentrionalis, were captured in mist nets in ... more Three adult male Northern Long-eared Bats, Myotis septentrionalis, were captured in mist nets in July 2004 in the LaBiche River Valley, southeastern Yukon. These are the first records of M. septentrionalis in the Yukon. Further survey work is needed to delineate the extent of the range and population structure of this and other species of bats in northwestern North America.

Research paper thumbnail of Are the western water shrew (Sorex navigator) and American water shrew (Sorex palustris) morphologically distinct?

Canadian Journal of Zoology

Recent phylogenetic studies applying the mitochondrial cytochrome b (Cytb) gene and nuclear genes... more Recent phylogenetic studies applying the mitochondrial cytochrome b (Cytb) gene and nuclear genes revealed divergent cordilleran and boreal lineages in western populations of the Sorex palustris complex that are now recognized as distinct species: Sorex navigator (Baird, 1858) (western water shrew) and Sorex palustris Richardson, 1828 (American water shrew). Their distributions in western Canada are parapatric with a potential contact zone in the foothills of the Rocky Mountains. Using 80 specimens including 69 of known genotype, we assessed morphological differentiation in metric and nonmetric traits among S. navigator, S. palustris, and Sorex bendirii (Merriam, 1894) (marsh shrew), a sister species of S. navigator. Multivariate analyses revealed discrete morpho groups concordant with their genetic lineages and species-level divergence. Three individuals from northwestern British Columbia and one from southwestern Alberta with discordant mitochondrial and nuclear DNA aligned with t...

Research paper thumbnail of New records of the Ogilvie Mountains Collared Lemming (<em>Dicrostonyx nunatakensis</em>) in central Yukon

The Canadian Field-Naturalist, 2014

The Ogilvie Mountain Collared Lemming (Dicrostonyx nunatakensis Youngman, 1967), reported only fr... more The Ogilvie Mountain Collared Lemming (Dicrostonyx nunatakensis Youngman, 1967), reported only from the Ogilvie Mountains of central Yukon, is among the least known mammals in Canada. It was first discovered in 1961 and, since then, only 13 specimens had been collected, all from one mountain, in central Yukon. We conducted a targeted survey to determine the distribution of the species by trapping areas of apparently suitable habitat on 12 mountains within 40 km of the known location. Many of our traps were disabled by other mammals; however, we captured three Ogilvie Mountain Collared Lemmings on two mountains 25.9 km and 29.6 km from the original location. Our findings suggest that this lemming may be more widely distributed than indicated by earlier specimens. We suggest further surveys to delineate the range of the Ogilvie Mountain Collared Lemming.

Research paper thumbnail of A Tribute to Ian McTaggart-Cowan, 1910-2010, O.C., O.B.C., PhD, LL.D, F.R.S.C

The Canadian Field-Naturalist, 2010

Research paper thumbnail of First Records of the Southern Red-backed Vole, <em>Myodes gapperi</em>, in the Yukon

The Canadian Field-Naturalist, 2006

Twenty Southern Red-backed Voles, Myodes gapperi, were collected in July 2004 in the LaBiche Rive... more Twenty Southern Red-backed Voles, Myodes gapperi, were collected in July 2004 in the LaBiche River valley of southeastern Yukon. Specimens were identified using morphological characteristics and analysis of cytochrome b gene sequences. These are the first records of this species in the Yukon. No Northern Red-backed Voles, M. rutilus, were collected and it is not known whether the two species are sympatric or parapatric in the Yukon. Further survey work is needed in southeastern Yukon to better delineate the extent of the northwestern range of this species and the extent, if any, of introgression with M. rutilus.

Research paper thumbnail of A morphometric study of geographic variation in the snowshoe hare (Lepus americanus)

Canadian Journal of Zoology, Feb 14, 2011

Geographic variation in Lepus americanus was studied by multivariate analyses of 15 cranial measu... more Geographic variation in Lepus americanus was studied by multivariate analyses of 15 cranial measurements. A total of 1494 specimens from the entire geographic range were grouped into 37 geographic samples of males and females. Principal component analyses demonstrated that hares are largest in eastern North America, Alaska, and northwestern Canada, and smallest in the Pacific Northwest. Size clines exist in the Appalachian Mountains and western North America but size is relatively uniform throughout central Canada and the Great Lakes. A multiple regression of size with 16 climatic variables factors demonstrated that size and climate are strongly correlated; the size trends may reflect environmental selection. The differentiation of hares from the western Cordillera and Pacific coast, the similarity of populations from central Canada and the Great Lakes, and the clines in the Appalachians and western North America were evident in discriminant analyses. These patterns of variation among populations can be attributed to both gene flow and local selection. There is no evidence from the morphometric analyses for classifying populations into the 15 subspecies currently recognized.