Modeling Caribou Movements: Seasonal Ranges and Migration Routes of the Central Arctic Herd

Migration is an important component of the life history of many animals, but persistence of large-scale terrestrial migrations is being challenged by environmental changes that fragment habitats and create obstacles to animal movements. In northern Alaska, the Central Arctic herd (CAH) of barren-ground caribou (Rangifer tarandus granti) is known to migrate over large distances, but the herd’s seasonal distributions and migratory movements are not well documented. From 2003–2007, we used GPS radio-collars to determine seasonal ranges and migration routes of 54 female caribou from the CAH. We calculated Brownian bridges to model fall and spring migrations for each year and used the mean of these over all 4 years to identify areas that were used repeatedly. Annual estimates of sizes of seasonal ranges determined by 90% fixed kernel utilization distributions were similar between summer and winter ($\bar{X}$ = 27,929 SE = 1,064 and $\bar{X}$ = 26,585 SE = 4912 km², respectively). Overlap between consecutive summer and winter ranges varied from 3.3–18.3%. Percent overlap between summer ranges used during consecutive years ($\bar{X}$ = 62.4% SE = 3.7%) was higher than for winter ranges ($\bar{X}$ = 42.8% SE = 5.9%). Caribou used multiple migration routes each year, but some areas were used by caribou during all years, suggesting that these areas should be managed to allow for continued utilization by caribou. Restoring migration routes after they have been disturbed or fragmented is challenging. However, prior knowledge of movements and threats may facilitate maintenance of migratory paths and seasonal ranges necessary for long-term persistence of migratory species.     

New Insights into Non-Avian Dinosaur Reproduction and Their Evolutionary and Ecological Implications: Linking Fossil Evidence to Allometries of Extant Close Relatives

It has been hypothesized that a high reproductive output contributes to the unique gigantism in large dinosaur taxa. In order to infer more information on dinosaur reproduction, we established allometries between body mass and different reproductive traits (egg mass, clutch mass, annual clutch mass) for extant phylogenetic brackets (birds, crocodiles and tortoises) of extinct non-avian dinosaurs. Allometries were applied to nine non-avian dinosaur taxa (theropods, hadrosaurs, and sauropodomorphs) for which fossil estimates on relevant traits are currently available. We found that the reproductive traits of most dinosaurs conformed to similar-sized or scaled-up extant reptiles or birds. The reproductive traits of theropods, which are considered more bird-like, were indeed consistent with birds, while the traits of sauropodomorphs conformed better to reptiles. Reproductive traits of hadrosaurs corresponded to both reptiles and birds. Excluding Massospondylus carinatus , all dinosaurs studied had an intermediary egg to body mass relationship to reptiles and birds. In contrast, dinosaur clutch masses fitted with either the masses predicted from allometries of birds (theropods) or to the masses of reptiles (all other taxa). Theropods studied had probably one clutch per year. For sauropodomorphs and hadrosaurs, more than one clutch per year was predicted. Contrary to current hypotheses, large dinosaurs did not have exceptionally high annual egg numbers (AEN). Independent of the extant model, the estimated dinosaur AEN did not exceed 850 eggs (75,000 kg sauropod) for any of the taxa studied. This estimated maximum is probably an overestimation due to unrealistic assumptions. According to most AEN estimations, the dinosaurs studied laid less than 200 eggs per year. Only some AEN estimates obtained for medium to large sized sauropods were higher (200-400 eggs). Our results provide new (testable) hypotheses, especially for reproductive traits that are insufficiently documented or lacking from the fossil record. This contributes to the understanding of their evolution.     

Caribou movement as a correlated random walk

Movement is a primary mechanism coupling animals to their environment, yet there exists little empirical analysis to test our theoretical knowledge of this basic process. We used correlated random walk (CRW) models and satellite telemetry to investigate long-distance movements of caribou, the most vagile, non-volant terrestrial vertebrate in the world. Individual paths of migratory and sedentary female caribou were quantified using measures of mean move length and angle, and net squared displacements at each successive move were compared to predictions from the models. Movements were modelled at two temporal scales. For paths recorded through one annual cycle, the CRW model overpredicted net displacement of caribou through time. For paths recorded over shorter intervals delineated by seasonal behavioural changes of caribou, there was excellent correspondence between model predictions and observations for most periods for both migratory and sedentary caribou. On the smallest temporal scale, a CRW model significantly overpredicted displacements of migratory caribou during 3 months following calving; this was also the case for sedentary caribou in late summer, and in late winter. In all cases of overprediction there was significant positive autocorrelation in turn direction, indicating that movements were more tortuous than expected. In one case of underprediction, significant negative autocorrelation of sequential turn direction was evident, indicating that migratory caribou moved in straightened paths during spring migration to calving grounds. Results are discussed in light of known migration patterns and possible limiting factors for caribou, and indicate the applicability of CRW models to animal movement at vast spatial and temporal scales, thus assisting in future development of more sophisticated models of population spread and redistribution for vertebrates. Received: 14 July 1999 / Accepted: 15 November 1999     

Seasonal patterns of spatial fidelity and temporal consistency in the distribution and movements of a migratory ungulate

How animals use their range can have physiological, ecological, and demographic repercussions, as well as impact management decisions, species conservation, and human society. Fidelity, the predictable return to certain places, can improve fitness if it is associated with high‐quality habitat or helps enable individuals to locate heterogenous patches of higher‐quality habitat within a lower‐quality habitat matrix. Our goal was to quantify patterns of fidelity at different spatial scales to better understand the relative plasticity of habitat use of a vital subsistence species that undergoes long‐distance migrations. We analyzed a decade (2010–2019) of GPS data from 240 adult, female Western Arctic Herd (WAH) caribou (Rangifer tarandus) from northwest Alaska, U.S.A. We assessed fidelity at 2 spatial scales: to site‐specific locations within seasonal ranges and to regions within the herd''s entire range by using 2 different null datasets. We assessed both area and consistency of use during 6 different seasons of the year. We also assessed the temporal consistency of migration and calving events. At the scale of the overall range, we found that caribou fidelity was greatest during the calving and insect relief (early summer) seasons, where the herd tended to maximally aggregate in the smallest area, and lowest in winter when the seasonal range is largest. However, even in seasons with lower fidelity, we found that caribou still showed fidelity to certain regions within the herd''s range. Within those seasonal ranges, however, there was little individual site‐specific fidelity from year to year, with the exception of summer periods. Temporally, we found that over 90% of caribou gave birth within 7 days of the day they gave birth the previous year. This revealed fairly high temporal consistency, especially given the spatial and temporal variability of spring migration. Fall migration exhibited greater temporal variability than spring migration. Our results support the hypothesis that higher fidelity to seasonal ranges is related to greater environmental and resource predictability. Interestingly, this fidelity was stronger at larger scales and at the population level. Almost the entire herd would seek out these areas with predictable resources, and then, individuals would vary their use, likely in response to annually varying conditions. During seasons with lower presumed spatial and/or temporal predictability of resources, population‐level fidelity was lower but individual fidelity was higher. The herd would be more spread out during the seasons of low‐resource predictability, leading to lower fidelity at the scale of their entire range, but individuals could be closer to locations they used the previous year, leading to greater individual fidelity, perhaps resulting from memory of a successful outcome the previous year. Our results also suggest that fidelity in 1 season is related to fidelity in the subsequent season. We hypothesize that some differences in patterns of range fidelity may be driven by seasonal differences in group size, degree of sociality, and/or density‐dependent factors. Climate change may affect resource predictability and, thus, the spatial fidelity and temporal consistency of use of animals to certain seasonal ranges.     

Reconstruction of caribou evolutionary history in Western North America and its implications for conservation

The role of Beringia as a refugium and route for trans-continental exchange of fauna during glacial cycles of the past 2million years are well documented; less apparent is its contribution as a significant reservoir of genetic diversity. Using mitochondrial DNA sequences and 14 microsatellite loci, we investigate the phylogeographic history of caribou (Rangifer tarandus) in western North America. Patterns of genetic diversity reveal two distinct groups of caribou. Caribou classified as a Northern group, of Beringian origin, exhibited greater number and variability in mtDNA haplotypes compared to a Southern group originating from refugia south of glacial ice. Results indicate that subspecies R. t. granti of Alaska and R. t. groenlandicus of northern Canada do not constitute distinguishable units at mtDNA or microsatellites, belying their current status as separate subspecies. Additionally, the Northern Mountain ecotype of woodland caribou (presently R. t. caribou) has closer kinship to caribou classified as granti or groenlandicus. Comparisons of mtDNA and microsatellite data suggest that behavioural and ecological specialization is a more recently derived life history characteristic. Notably, microsatellite differentiation among Southern herds is significantly greater, most likely as a result of human-induced landscape fragmentation and genetic drift due to smaller population sizes. These results not only provide important insight into the evolutionary history of northern species such as caribou, but also are important indicators for managers evaluating conservation measures for this threatened species.     

Trade-off between reproduction and moult – a comparison of three Fennoscandian pied flycatcher populations

Organisms that reproduce at high latitudes are assumed to have evolved several adaptations to the short summer. For birds, and especially for long-distance migrants, there is a time constraint because both reproduction and moult must be completed before autumn migration. It has therefore been assumed that birds at northern latitudes must initiate their moult during reproduction more often than birds at low latitudes. To investigate how passerine birds breeding at different latitudes allocate their time between reproduction and moult, we compared timing of these activities during three consecutive breeding seasons in three widely separated populations of the pied flycatcher Ficedula hypoleuca. Our results show that the frequency of individuals with moult-breeding overlap, and moult initiation in relation to breeding stage, varied considerably among populations and years. In all three populations, female moult initiation was restricted to the late nestling period. The males had a more pronounced moult-breeding overlap than the females, but its duration was similar in all three study areas. Thus, there was no evidence for a more pronounced moult-breeding overlap at high compared with low latitudes. These results suggest that pied flycatchers sometimes accept a moult-breeding overlap, but that the time gained by having too extensive an overlap between reproduction and moult does not outweigh the associated costs. Long-distance migrants breeding at northern latitudes apparently experience a trade-off between reproduction and somatic investment during moult. We therefore suggest that a pronounced moult-breeding overlap is not a typical strategy used by long-distance migrants to adjust to the short breeding season at northern latitudes. Received: 7 May 1998 / Accepted: 24 August 1998     

Wolves,white‐tailed deer,and beaver: implications of seasonal prey switching for woodland caribou declines

Population increases of primary prey can negatively impact alternate prey populations via demographic and behavioural responses of a shared predator through apparent competition. Seasonal variation in prey selection patterns by predators also can affect secondary and incidental prey by reducing spatial separation. Global warming and landscape changes in Alberta's bitumen sands have resulted in prey enrichment, which is changing the large mammal predator–prey system and causing declines in woodland caribou Rangifer tarandus caribou populations. We assessed seasonal patterns of prey use and spatial selection by wolves Canis lupus in two woodland caribou ranges in northeastern Alberta, Canada, that have undergone prey enrichment following recent white‐tailed deer Odocoileus virginianus invasion. We determined whether risk of predation for caribou (incidental prey) and the proportion of wolf‐caused‐caribou mortalities varied with season. We found that wolves showed seasonal variation in primary prey use, with deer and beaver Castor canadensis being the most common prey items in wolf diet in winter and summer, respectively. These seasonal dietary patterns were reflected in seasonal wolf spatial resource selection and resulted in contrasting spatial relationships between wolves and caribou. During winter, wolf selection for areas used by deer maintained strong spatial separation between wolves and caribou, whereas wolf selection for areas used by beaver in summer increased the overlap with caribou. Changing patterns in wolf resource selection were reflected by caribou mortality patterns, with 76.2% of 42 adult female caribou mortalities occurring in summer. Understanding seasonal patterns of predation following prey enrichment in a multiprey system is essential when assessing the effect of predation on an incidental prey species. Our results support the conclusion that wolves are proximately responsible for woodland caribou population declines throughout much of their range.     

Assessment of Canada Lynx Research and Conservation Needs in the Southern Range: Another Kick at the Cat

Abstract The ecology of Canada lynx (Lynx canadensis) and their main prey, snowshoe hares (Lepus americanus), is poorly understood in southern Canada and the contiguous United States compared to the boreal forest of Canada and Alaska, USA, where both species are well studied. However, given recent listing of lynx under the Endangered Species Act, accurate understanding of lynx and snowshoe hare ecology and conservation requirements in the United States is a high priority. We critically examined unchallenged perceptions and important research needs related to lynx and hare ecology and conservation at the southern extent of their range. Contrary to popular dogma, lynx do not require old-growth forest for denning, but further research on lynx and hare use of fragmented landscapes at lower latitudes is required. The contention that southern lynx are subject to higher interference or exploitative competition compared to their northern counterparts remains without strong empirical support. Lynx rely more on red squirrels (Tamiasciurus hudsonicus) and possibly other alternate prey at lower latitudes, but hares are the predominant food type for lynx across their range. Southern lynx and hare populations do not exhibit periodic cyclicity, but harvest statistics suggest that lynx abundance in the southern range is highly variable, implying that numerical fluctuations likely are fueled by immigration from Canada. Southern lynx population viability in the absence of ingress is suspect and thus maintaining connectivity with northern areas of occupancy should be a priority. Successful conservation of lynx populations in the contiguous United States will require 1) improved understanding of lynx population and habitat ecology at lower latitudes, 2) protection and management of large tracts of lynx and snowshoe hare habitat, and 3) ensured connectivity between lynx populations at the core and periphery of the species' range. However, in light of the numerous challenges facing conservation of populations of many species at their southern distributional limit, the long-term prognosis for lynx in the southern range currently is uncertain.     

Ecological factors associated with the breeding and migratory phenology of high-latitude breeding western sandpipers

Environmental conditions influence the breeding and migratory patterns of many avian species and may have particularly dramatic effects on long-distance migrants that breed at northern latitudes. Environment, however, is only one of the ecological variables affecting avian phenology, and recent work shows that migration tactics may be strongly affected by changes in predator populations. We used long-term data from 1978 to 2000 to examine the interactions between snowmelt in western Alaska in relation to the breeding or migration phenologies of small shorebirds and their raptor predators. Although the sandpipers’ time of arrival at Alaskan breeding sites corresponded with mean snowmelt, late snowmelts did delay breeding. These delays, however, did not persist to southward migration through British Columbia, likely due to the birds’ ability to compensate for variance in the length of the breeding season. Raptor phenology at an early stopover site in British Columbia was strongly related to snowmelt, so that in years of early snowmelt falcons appeared earlier during the sandpipers’ southbound migration. These differential effects indicate that earlier snowmelt due to climate change may alter the ecological dynamics of the predator–prey system.     

Caribou Distribution and Movements in a Northern Alaska Oilfield

As industrial development increases in the range of barren-ground caribou (Rangifer tarandus granti) across the warming Arctic, the need to understand the responses of caribou to development and to assess the effectiveness of mitigation measures increase accordingly. The Central Arctic Herd (CAH) of caribou ranges across northern Alaska, USA, and the herd's summer range includes the Prudhoe Bay and Kuparuk oilfields, where the herd has been exposed to oil development for >4 decades. We used location data from global positioning system (GPS) radio-collars deployed on female CAH caribou for 106 collar-years, recording locations every 2 hours during 2008–2019, to examine caribou distribution and movements during 7 different seasons of the year in relation to infrastructure in the Kuparuk oilfield, which is characterized by more design improvements and mitigation measures than the older Prudhoe Bay oilfield. We examined movement metrics in terms of distance to gravel infrastructure (roads and pads) and time before and after movements across infrastructure (crossings). We also employed integrated step-selection analysis to compare caribou movements with random movements. Caribou distribution was influenced by insect activity, distance to coast, landcover, and terrain ruggedness, and we found large seasonal differences in caribou responses to infrastructure. Consistent with previous research findings, avoidance of areas near roads and pads was strongest during the calving season and some caribou used roads and pads as insect-relief habitat when oestrid flies (warble fly [Hypoderma tarandi] and nose bot fly [Cephenemyia trompe]) were active. Caribou moved through the Kuparuk oilfield repeatedly during summer, averaging >2 road or pad crossings a day when harassment by mosquitoes (Aedes [Ochlerotatus] spp.) and oestrid flies were the predominant factors influencing caribou movements. Caribou moved faster while crossing roads and pads but showed little pattern in speed or turn angle with distance to roads and pads. These results demonstrate that the effects of petroleum development on a caribou herd with long-term exposure to industrial activity vary widely by season. Maternal caribou avoid active roads and pads during calving, but the incorporation of appropriate mitigation measures in oilfield design allows caribou to move through the Kuparuk oilfield during other snow-free seasons. © 2020 The Wildlife Society.     

High latitudes and high genetic diversity: phylogeography of a widespread boreal bird, the gray jay (Perisoreus canadensis)

We describe range-wide phylogeographic variation in gray jays (Perisoreus canadensis), a boreal Nearctic corvid that occurs today primarily in recently glaciated regions. Phylogenetic analysis of mitochondrial DNA (1041 base pairs ND2 gene; N=205, 50 localities) revealed four reciprocally monophyletic groups. One widespread clade occurs across the North American boreal zone, from Newfoundland to Alaska and southwest into Utah. Three other clades occur at lower latitudes in the montane West in Colorado, the northern Rocky Mountains, and the Pacific Northwest respectively. The geographic distribution of clades in gray jays corresponds with a general pattern that is emerging for boreal taxa, having one widespread northern clade and one or more geographically restricted southwestern clades. Population genetic analyses indicate that the larger boreal clade is genetically structured and harbors significantly more genetic diversity than those clades occurring at lower latitudes. Species distribution modeling (SDM) revealed multiple putative Pleistocene refugia including several occurring at higher latitudes. We suggest that multiple post-glacial colonization routes, some of which originate from these northern refugia, are responsible for the relatively high genetic diversity at high latitudes. Conversely, lower latitude clades show little variation, probably as a result of historical restriction to smaller geographical areas with smaller long-term population sizes. This 'upside-down' pattern of genetic diversity contrasts with the conventional view that populations of north-temperate species occupying previously glaciated habitats should possess lower levels of diversity than their southern counterparts.     

Intraspecific alkaloid variation in ladybird eggs and its effects on con- and hetero-specific intraguild predators

Sexual size dimorphism (SSD) is a common phenomenon in animals. In many species females are substantially larger than males. Because body size plays a central role in modulating the body temperature (T _b) of ectotherms, intersexual differences in body size may lead to important intersexual differences in thermoregulation. In addition, because SSD is realized by differences in growth rate and because growth rate is strongly temperature dependent in ectotherms, a conflict between male reproductive behaviour and thermoregulation may affect the expression of SSD. In this study, we investigated the thermal implications of SSD in a reptile exhibiting spectacular female-biased SSD: the northern map turtle (Graptemys geographica). Over three seasons, we collected >150,000 measurements of T _b in free-ranging adult and juvenile northern map turtles using surgically implanted miniature temperature loggers. Northern map turtles exhibited seasonal patterns of thermoregulation typical of reptiles in northern latitudes, but we found that large adult females experienced a lower daily maximum T _b and a narrower daily range of T _b than adult males and small juvenile females. In addition, despite more time spent basking, large adult females were not able to thermoregulate as accurately as small turtles. Our findings strongly suggest that body size limits the ability to thermoregulate accurately in large females. By comparing thermoregulatory patterns between adult males and juvenile females of similar body size, we found no evidence that male reproductive behaviours are an impediment to thermoregulation. We also quantified the thermal significance of basking behaviour. We found, contrary to previous findings, that aerial basking allows northern map turtles to raise their T _b substantially above water temperature, indicating that basking behaviour likely plays an important role in thermoregulation.     

Caribou Use of Habitat Near Energy Development in Arctic Alaska

Increasing demands for energy have generated interest in expanding oil and gas production on the North Slope of Alaska, USA, raising questions about the resilience of barren-ground caribou (Rangifer tarandus) populations to new development. Although the amount of habitat lost directly to energy development in the Arctic will likely be relatively small, there are significant concerns about habitat that may be indirectly affected because of caribou avoidance behaviors. Behavioral responses to energy development for wildlife have been documented, but such responses are often assumed to dissipate over time, despite scant information on the ability of animals to habituate. To understand the long-term effects of energy development on barren-ground caribou, we investigated the behavior of the Central Arctic Herd in northern Alaska, which has been exposed to oil development on its summer range for approximately 40 years. Using recent (2015–2017) location data from global positioning system (GPS)-collared females, we conducted a zone of influence analysis to assess whether caribou reduced their use of habitat near energy development, and if so, the distance the effects attenuated. We conducted this analysis for the calving, post-calving, and mosquito harassment periods when caribou exhibit distinct resource selection patterns, and contrasted our results to past research that investigated the responses of the Central Arctic Herd immediately following the construction of the oil fields. Despite the long-term presence of energy development within the Central Arctic Herd summer range, we found that female caribou exhibited avoidance responses to infrastructure during all time periods, although the effects waned across the summer. Caribou reduced their use of habitat within 5 km of development during the calving period, within 2 km during the post-calving period, and within 1 km during the mosquito harassment period; these areas were predicted to overlap 12%, 15%, and 17% of important calving, post-calving, and mosquito period habitat, respectively. During the calving period, the indirect effects we observed were similar to those observed in past research, whereas during the post-calving and mosquito periods, we detected avoidance responses that had not been previously reported. These findings corroborate a growing body of evidence suggesting that habituation to industrial development in caribou in the Arctic is likely to be weak or absent, and emphasizes the value of minimizing the footprint of infrastructure within important seasonal habitat to reduce behavioral effects to barren-ground caribou. © 2019 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Age-related timing of Short-toed Snake Eagle Circaetus gallicus migration along detoured and direct flyways

Capsule: Juvenile Short-toed Snake Eagles Circaetus gallicus hatching in the peripheral populations of Greece and Italy have limited opportunities for social learning of migration routes compared to those hatched elsewhere.

Aims: To test the prediction that there would be a higher degree of migration synchrony between adult and juvenile Short-toed Snake Eagles originating from peripheral populations and using an extremely detoured flyway, when compared to other populations using a direct overland flyway.

Methods: We use linear regression models to compare seasonal changes in the age distribution of migrating Short-toed Snake Eagles counted at two migration watch-sites in Italy (Arenzano) and Georgia (Batumi), along a detoured and a direct flyway, respectively.

Results: Juveniles migrated a few days later than adults at both sites and the age ratios recorded at these two sites was similar. The daily proportion of juveniles increased along a similar slope during the migration season, thus showing a similar degree of synchrony between the age classes on both flyways.

Conclusions: Contrary to our hypothesis, juvenile and adult migration is not more synchronized in peripheral populations using a detoured flyway compared to a core population using a direct migration flyway. Our results suggest that juveniles do not learn detours to complete trans-Mediterranean migration from their parents, but from other elders.     

Differences in Tooth Microwear of Populations of Caribou (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>, Ruminantia,Mammalia) and Implications to Ecology,Migration, Glaciations and Dental Evolution

Tooth microwear was analyzed for a large sample of wild-shot barren-ground caribou (Rangifer tarandus groenlandicus) from the Kaminuriak population of eastern Canada. This sample was compared to the microwear of specimens from three Pleistocene localities in North America (Alaska) and western Europe (Caune de l’Arago in France and Salzgitter in Germany). The results show that the extant samples from eastern Canada have seasonal variation in microwear and presumably in diet. The differences in microwear between the various seasons may reflect a cyclic migration of the population within a year. The extinct population from Alaska has extremely blunt teeth (mesowear), as blunt as those of modern zebras and bison. This observation is corroborated by the lowest number of microwear pits. The findings are untypical, as most typical caribou teeth have sharper apices, and we interpret this as an indication of a local habitat that was different with animals feeding on non-typical vegetation. The combination of Rangifer from Caune de l’Arago and Salzgitter reveals a pattern in microwear variability. The Salzgitter is interglacial and shows a greater diversity of browsing (broad spectrum on average number of pits) than the glacial Caune de l’Arago. The interglacial population from Salzgitter is interesting because it shows several different types of browsing. Collectively all the Rangifer teeth show that diet of a brachydont taxon can vary across most of the dietary morphospace of ungulates as represented by tooth microwear. The three Pleistocene samples exhibit microwear that is different from the extant population in question. This observation implies that the recent diet of Rangifer has changed from the typical caribou diet in the past. This indicates dietary change within a species. This is important because it represents dietary evolution without changes in tooth morphology.     

Lack of sperm production and sperm storage by arctic-nesting shorebirds during spring migration

Birds nesting at high latitudes may copulate during migration to arrive on the nesting grounds ready to breed. We surveyed 12 species of shorebirds during spring migration to determine whether (1) males produced abundant sperm and (2) females harboured functional sperm storage tubules (SSTs). Sperm production by males on migration was rare. Only four of seven species (9.8% of 41 males) of long-distance migrants harboured sperm, whereas all four species (100% of eight males) of short-distance migrants held sperm. In females, no long-distance migrants held sperm in their SSTs ( n  = 28 females) and SSTs were small compared to long-distance migrants collected on their breeding grounds. Our results indicate that shorebirds nesting at high latitudes were generally not reproductively active during migration and that any sexual behaviour on migration is unlikely to lead directly to fertilizations.     

Theropod fauna from southern Australia indicates high polar diversity and climate-driven dinosaur provinciality

The Early Cretaceous fauna of Victoria, Australia, provides unique data on the composition of high latitude southern hemisphere dinosaurs. We describe and review theropod dinosaur postcranial remains from the Aptian-Albian Otway and Strzelecki groups, based on at least 37 isolated bones, and more than 90 teeth from the Flat Rocks locality. Several specimens of medium- and large-bodied individuals (estimated up to ~8.5 metres long) represent allosauroids. Tyrannosauroids are represented by elements indicating medium body sizes (~3 metres long), likely including the holotype femur of Timimus hermani, and a single cervical vertebra represents a juvenile spinosaurid. Single specimens representing medium- and small-bodied theropods may be referrable to Ceratosauria, Ornithomimosauria, a basal coelurosaur, and at least three taxa within Maniraptora. Thus, nine theropod taxa may have been present. Alternatively, four distinct dorsal vertebrae indicate a minimum of four taxa. However, because most taxa are known from single bones, it is likely that small-bodied theropod diversity remains underestimated. The high abundance of allosauroids and basal coelurosaurs (including tyrannosauroids and possibly ornithomimosaurs), and the relative rarity of ceratosaurs, is strikingly dissimilar to penecontemporaneous dinosaur faunas of Africa and South America, which represent an arid, lower-latitude biome. Similarities between dinosaur faunas of Victoria and the northern continents concern the proportional representatation of higher clades, and may result from the prevailing temperate-polar climate of Australia, especially at high latitudes in Victoria, which is similar to the predominant warm-temperate climate of Laurasia, but distinct from the arid climate zone that covered extensive areas of Gondwana. Most dinosaur groups probably attained a near-cosmopolitan distribution in the Jurassic, prior to fragmentation of the Pangaean supercontinent, and some aspects of the hallmark 'Gondwanan' fauna of South America and Africa may therefore reflect climate-driven provinciality, not vicariant evolution driven by continental fragmentation. However, vicariance may still be detected at lower phylogenetic levels.     

Modeling the Western Arctic Caribou Herd During a Positive Growth Phase: Potential Effects of Wolves and Radiocollars

ABSTRACT Population modeling exercises can lead to both expected and unexpected results useful for wildlife research and management, even though inferences must often be qualitative, given underlying assumptions. Our main objective was to use empirical data on wolf (Canis lupus) kill rates and growth of the Western Arctic caribou (Rangifer tarandus) herd (WAH) of Alaska, USA, to assess the potential for predator regulation. We used available data and published literature to construct a deterministic density-dependent population model fitted to trends of the WAH from 1976 through 2003. By increasing wolf densities in the baseline model, we failed to reject the hypothesis that wolves at a density of 6.5 wolves per 1,000 km² could regulate a caribou herd at a density of 0.4 caribou per km². In addition, our model may be conservative by underestimating the regulatory potential of wolves. We suggest that this relatively simple predator-preysystem shows signs of a predation—food 2-state model. Elasticities from matrix models may be deceiving. Although herd growth is most sensitive to changes in adult female survival, survival of younger cohorts may be more easily influenced by natural conditions or management action. Management of the WAH near maximum sustained yield may not be attainable if desired, but modeling exercises such as this elucidate options. In conducting this research, we also discovered by Monte Carlo simulation that survival and productivity data from radiocollared females and calves were negatively biased and failed to predict herd growth. Thus, researchers should consider potential effects of neck collars on vital rates of female tundra caribou and concomitant offspring when using sample data to model population dynamics or test hypotheses.     

Isotopic tracking of foraging and long-distance migration in northeastern Pacific pinnipeds

We investigated the impact of foraging location (nearshore vs offshore) and foraging latitude (high vs middle) on the carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope compositions of bone collagen of northern fur seals (Callorhinus ursinus), harbor seals (Phoca vitulina), California sea lions (Zalophus californianus), and northern elephant seals (Mirounga angustirostris). Nearshore-foraging harbor seals from California had δ¹³C values 2.0‰ higher than female northern elephant seals foraging offshore at similar latitudes. Likewise, nearshore-foraging harbor seals from Alaska had values 1.7‰ higher than male northern fur seals, which forage offshore at high latitudes. Middle-latitude pinnipeds foraging in either the nearshore or offshore were ¹³C enriched by ∼1.0‰ over similar populations from high latitudes. Male northern elephant seals migrate between middle and high latitudes, but they had δ¹³C values similar to high-latitude, nearshore foragers. Female northern fur seal δ¹³C values were intermediate between those of high- and middle-latitude offshore foragers, reflecting their migration between high- and middle-latitude waters. The δ¹³C values of California sea lions were intermediate between nearshore- and offshore-foraging pinnipeds at middle latitudes, yet there was no observational support for the suggestion that they use offshore food webs. We suggest that their “intermediate” values reflect migration between highly productive and less-productive, nearshore ecosystems on the Pacific coasts of California and Mexico. The relative uniformity among all of these pinnipeds in δ¹⁵N values, which are strongly sensitive to trophic level, reveals that the carbon isotope patterns result from differences in the δ¹³C of organic carbon at the base of the food web, rather than differences in trophic structure, among these regions. Finally, the magnitude and direction of the observed nearshore-offshore and high-to middle-latitude differences in δ¹³C values suggest that these gradients may chiefly reflect differences in rates and magnitudes of phytoplankton production as well as the δ¹³C value of inorganic carbon available for photosynthesis, rather than the input of ¹³C-enriched macroalgal carbon to nearshore food webs. Received: 8 September 1998 / Accepted: 24 February 1999     

Longevity and growth rate estimates for a polar dinosaur: a Pachyrhinosaurus (Dinosauria: Neoceratopsia) specimen from the North Slope of Alaska showing a complete developmental record

Our knowledge of growth dynamics in large ceratopsian dinosaurs is very poor, in part, due to the paucity of quantifiable age markers such as growth lines in their bones. We sought marker-based, osteohistological evidence for ceratopsid age structure from high Arctic paleolatitudes based on the observations that: (1) extant mammals from high latitudes better express growth lines in their hard tissues than those from lower latitudes, and (2) the occurrence of accentuated growth banding in teeth from Arctic dinosaurs. We examined the long bones in the specimens of Pachyrhinosaurus sp. from the early Maastrichtian of northern Alaska, and found conspicuous osseous banding. Histological analysis of the spacing, structure, pattern and numbers of these bands in the femur of a very large specimen suggests that they are not a taphonomic artefact, rather they appear to reflect annual growth cycling. Counts and measurements of the growth zones suggest that the animal showed rapid linear growth early in ontogeny, sexual maturity in perhaps the 9th year of life, and that it died in the 19th year of life. Our data adds to a growing body of evidence that a genetically distinct northern dinosaurian fauna existed at high paleolatitudes in Alaska during the Late Cretaceous.