Eimeria dalli sp. n. (Protozoa: Eimeriidae) from Dall Sheep Ovis dalli

Eimeria dalli sp. n. is described from fecal samples collected from Dall sheep, Ovis dalli, from the Kenai Peninsula, Alaska. The oocysts are spherical or subspherical with mean dimensions of 43.7 × 37.4 μm. The outer oocyst wall is rough and irregular. No micropyle, micropylar cap or residuum was observed. Sporocysts were elongate ovoid with mean dimensions of 19.0 × 10.7 μm. Stieda bodies were not discernible.     

Cranial asymmetry in a Dall sheep ram (Ovis dalli dalli).

The horns of a 13-yr-old Dall sheep ram (Ovis dalli dalli) from the Sheep Mountain herd in Kluane National Park (Yukon, Canada) had unique characteristics. The right horn was 1,127 mm long, while the left horn was only 861 mm long and compressed and distorted at its base. The reduced growth of the left horn was due to chronic epidermitis and osteomyelitis of the cornual process, which began when the ram was 5 yr of age.     

Reproductive,behavioral, and endocrine characteristics of the Dall's sheep (Ovis dalli dalli)

Failed interspecific embryo transfer between Dall's sheep (Ovis dalli dalli) and domestic ewes (Ovis aries), and a paucity of physiological data available for the Dall's sheep, provided incentive for developing a reproductive database in Dall's sheep. Urine samples were collected on a regular basis from a captive herd of Dall's sheep during the breeding to lambing interval over a 3 year period. To provide comparative endocrine data during pregnancy, samples also were collected during a single gestation period from five Suffolk ewes. All urine samples were analyzed for pregnanediol-3-glucuronide (PdG) and estrone conjugates (EC). Behavioral observations from Dall's sheep were made each year during the rut and lambing periods to calculate estrous cycle and gestation lengths. Placentas from Dall's sheep (n = 20) and Suffolk ewes (n = 10) were collected, the total number of cotyledons counted, and the length, width, area, and weight of eight cotyledons/horn from each placenta calculated. Endocrine and behavioral results indicated that the Dall's sheep is seasonally polyestrous and monovulatory, with a mean estrous cycle length of 18.2 ± 0.3 days, a mean luteal phase of 11.8 ± 0.5 days, and an average gestation length of 171.6 ± 0.9 days. While hormonal patterns for pregnancy generally were similar between the two species, there was a pronounced difference in the magnitude of hormone concentrations, particularly PdG, and an absence of a marked preparturition EC rise in domestic sheep. Cotyledon numbers, weight, and area were smaller (P < 0.05) in the Dall's sheep, compared to Suffolk ewes. These data suggest that unsuccessful interspecies embryo transfer attempts may have been due to failed fetal/maternal communications resulting from hormonal and/or placental differences. © 1996 Wiley-Liss, Inc.     

An epizootic of Mycoplasma ovipneumoniae infection in captive Dall's sheep (Ovis dalli dalli)

In late spring of 1986, 10 of 23 Dall's sheep (Ovis dalli dalli) at the Metropolitan Toronto Zoo were moved to a new exhibit, where all developed severe respiratory signs refractory to anthelmintic and antibiotic therapy. In July, two animals died with chronic active bronch-pneumonia, and a third was euthanized because of pneumonia several months later. Bacteria were not isolated from the lungs of the first, steptococci and Pasteurella hemolytica were isolated from the other two, respectively; Mycoplasma ovipneumoniae was isolated from both. Pulmonary lesions in all three sheep were consistent with Mycoplasma sp. infection. Nasal swabs of the remaining animals yielded no consistent bacterial isolates; however, four of eight sheep were positive for M. ovipneumoniae. Viral cultures yielded an as yet unidentified herpesvirus. Sheep in the original and new herds had no serologic titers to parainfluenza-3, equine viral rhinopneumonitis, or infectious bovine rhinotracheitis, and had variable titers against bovine respiratory syncytial virus. No titers against M. ovipneumoniae were present in 13 sheep still in the original exhibit, but titers varied from 1:32 to 1:256 in eight pneumonic sheep. Sera taken from three sheep before or early in the outbreak were all negative for antibody to M. ovipneumoniae. Two of the affected Dall's sheep had been in contact with domestic sheep in the winter of 1985-1986, and M. ovipneumoniae was subsequently cultured from the domestic flock. Exposure to a new pathogen, and environmental and social stress in a new exhibit may have resulted in this severe disease in Dall's sheep.     

Parasites of the Dall's porpoise (Phocoenoides dalli True)

The prevalences of three helminths, Campula oblonga, Halocercus dalli and Crassicauda sp., recovered from Dall's porpoises which were net-entrapped incidentally in the vicinity of the Western Aleutian Islands in the northwest Pacific are reported. Specimens of Campula oblonga were found within the bile ducts of 46% of 127 livers examined. The prevalence of hepatic trematodiasis increased with the age of the host. Pulmonary nodules associated with Halocercus dalli were noted in 71% of the Dall's porpoises. Adult H. dalli were recovered from the main stem bronchi of heavily infected lungs. Younger animals exhibited a relatively higher prevalence. Specimens of Crassicauda sp. were found within the main lactiferous canal of 69% of 29 mammary glands examined. The prevalence was highest in mature porpoises. Possible detrimental effects and the modes of transmission of the three species of parasites are also considered.     

Mate Guarding in Male Dall's Porpoises (Phocoenoides dalli)

Mate guarding, whereby a male closely attends and defends a fertile female from extra‐pair matings, is one mating tactic males of many species use to protect their paternity. Although female defense occurs in many species of terrestrial mammal, comparable examples among cetaceans are largely absent, potentially as a result of the wide dispersion and mobility of females and their prey. Here, we investigate whether the close association of individual male Dall's porpoises with individual females during the breeding season is consistent with mate guarding. As mate guarding is predicted to be costly, and in other taxa is often associated with a reduction in foraging efficiency, we also examine whether males trade‐off this activity with time at depth. Males maintained longer associations and closer distances with female partners than with male ones. They also surfaced in greater synchrony with, and more often approached, their female partners than male ones. In contrast to males with male partners, males paired with females engaged in agonistic interactions with other adult males, and infrequently affiliated with extra‐pair individuals. These data suggest males are actively attempting to maintain their associations with females, while also acting to reduce female extra‐pair copulations and increase their own paternity. Guarding males also undertook shorter dives than non‐guarding males, suggesting that they trade‐off time at depth with guarding. Such a trade‐off is likely to involve a reduction in foraging opportunities, due to a decrease in time spent at foraging depth. Mate guarding in this species may be facilitated by the relatively smaller size and decreased mobility of newly calved, estrous females, particularly if females also benefit from guarding.     

Population genetic structure of North American thinhorn sheep (Ovis dalli)

The thinhorn sheep (Ovis dalli ssp.) provides a rare example of a North American large mammal that occupies most of its native range and maintains close to ancestral population size. There are currently two recognized subspecies, Dall's sheep (O. d. dalli) and Stone's sheep (O. d. stonei), the validity of which remains uncertain. We investigated the spatial genetic structure of thinhorn sheep populations representing both subspecies by genotyping individuals (n = 919) from across the species range at 12 variable microsatellite loci. We found high levels of genetic diversity within (HE = 0.722) and significant genetic structure among the 24 sampled areas (FST = 0.160). Genetic distance measures and Bayesian clustering analyses revealed the presence of at least eight subpopulations that are delineated by mountain range topology. A strong overall pattern of isolation-by-distance is evident across the sampling range (r = 0.75, P < 0.001) suggesting limited dispersal and extensive philopatry. Partial Mantel tests of this relationship showed mountain range distinctions represent significant barriers to gene flow (P = 0.0001), supporting the Bayesian analyses. Genetic structure was more strongly pronounced in southern Yukon and Alaska than elsewhere. We also show evidence for genetic differences between the two currently recognized thinhorn subspecies.     

Sex reversal in pairs of Lythrypnus dalli: behavioral and morphological changes

In Lythrypnus dalli, the bluebanded goby, reproductive success is primarily determined by functional sex, and functional sex is determined largely by rank in the dominance hierarchy. In most natural social groups of L. dalli, one male is at the apex of the hierarchy, and 1 to 7 females are lower in rank. When a male exits the group, a female ascends to the top of the hierarchy and becomes a male. We have examined this process in a simplified environment--a pair of females--that allows us to identify behavior associated with the formation of a dominance relationship and any other phenotypic changes associated with dominance, sex change or both. We found that pairs of L. dalli females quickly and readily form stable dominance relationships, with the dominant fish changing sex into a male. This dominant animal also rapidly increases in body size and length of its dorsal fin. In summary, dominant L. dalli females change sex in this simplified environment, providing excellent opportunities to examine the early behavioral and morphological changes associated with dominance and sex change.     

The phylogeographic pattern of mitochondrial DNA variation in the Dall's porpoise Phocoenoides dalli

We used 11 restriction endonucleases to study mtDNA variation in 101 Dall's porpoises Phocoenoides dalli from the Bering Sea and western North Pacific. There was little phylogeographic patterning among the 34 mtDNA haplotypes identified in this analysis, suggesting a strong historical connection among populations across this region. Nonetheless, mtDNA variation does not appear to be randomly distributed in this species. Both G_ST and AMOVA uncovered significant differences in the distribution of mtDNA variation between the Bering Sea and western North Pacific populations. These mtDNA results, coupled with differences in allozyme variation and parasite infestation, support the demographic distinctiveness of Bering Sea and western North Pacific stocks of Dall's porpoise. The lack of a strong phylogeographic orientation of mtDNA haplotypes within the Dall's porpoise is similar to the pattern reported in other vertebrates such as coyotes, blackbirds, chickadees, marine catfish, and catadromous eels. Like Dall's porpoise, these species are broadly distributed, and have large populations linked by moderate to high levels of gene flow. However, the more complex, deeply branched phylogenetic network of mtDNA haplotypes within Dall's porpoise, relative to these other vertebrates, suggests important differences between these species in the forces shaping mtDNA variation. One such force is the effective size of female populations, which appears to have been comparatively large and stable in Dall's porpoise.     

Protostrongylid parasites and pneumonia in captive and wild thinhorn sheep (Ovis dalli)

We describe health significance of protostrongylid parasites (Parelaphostrongylus odocoilei and Protostrongylus stilesi) and other respiratory pathogens in more than 50 naturally infected Dall's sheep (Ovis dalli dalli) from the Mackenzie Mountains, Northwest Territories (1998-2002) as well as in three Stone's sheep (O. d. stonei) experimentally infected with P. odocoilei (2000-2002). Histological lesions in the brain and distribution of P. odocoilei in the muscles of experimentally and naturally infected sheep were consistent with a previously hypothesized "central nervous system to muscle" pattern of migration for P. odocoilei. Dimensions of granulomas associated with eggs of P. odocoilei and density of protostrongylid eggs and larvae in the cranial lung correlated with intensity of larvae in feces, and all varied with season of collection. Prevalence of P. stilesi based on the presence of larvae in feces underestimated true prevalence (based on examination of lungs) in wild Dall's sheep collected in summer and fall. Similarly, counts of both types of protostrongylid larvae in feces were unreliable indicators of parasitic infection in wild Dall's sheep with concomitant bacterial pneumonia associated with Arcanobacterium pyogenes, Pasteurella sp., and Mannheimia sp. Diffuse, interstitial pneumonia due to P. odocoilei led to fatal pulmonary hemorrhage and edema after exertion in one experimentally infected Stone's sheep and one naturally infected Dall's sheep. Bacterial and verminous pneumonia associated with pathogens endemic in wild Dall's sheep in the Mackenzie Mountains caused sporadic mortalities. There was no evidence of respiratory viruses or bacterial strains associated with domestic ruminants, from which this population of wild sheep has been historically isolated.     

Elevated 11-ketotestosterone during paternal behavior in the Bluebanded goby (Lythrypnus dalli)

The relationship between androgens and paternal behavior is not straightforward, potentially because of the diversity of tasks a male must undertake to maximize reproductive success, notably alternating between courtship, aggression, and offspring care. In some species, these events are separated in time, but in others they are coincident. The endocrine profiles of species that simultaneously court, parent, and defend a nest, such as male bluebanded gobies (Lythrypnus dalli), are not well understood. We sampled a potent fish androgen, 11-ketotestosterone (KT), at different life history stages (experienced parenting males, experienced males not actively parenting, inexperienced males with their first clutch, and females), to examine this relationship. We found that experienced parenting L. dalli males have the highest KT levels of any group, while none of the other groups differed significantly. Males showed elevated KT levels when they had eggs compared to when they did not. Our data suggest that KT facilitates at least some aspects of parental care in L. dalli.     

Social status determines sexual phenotype in the bi-directional sex changing bluebanded goby Lythrypnus dalli

The behavioural mechanisms and patterns of protandrous sex change in bluebanded gobies Lythrypnus dalli were investigated and compared to the well-described behaviour patterns of protogynous sex change. To do this, unisex groups of males and females were established; behavioural and anatomical changes were recorded over a 42 day period as social status and sexual phenotype were determined. In all cases, social status, rather than the expression of a particular behaviour, accurately predicted final sexual phenotype. Rates of submissive behaviour, but not aggressive behaviour, were predictive of each discrete status class. Multiple individuals changed sex simultaneously if their sexual phenotype and social status were discordant, a novel finding suggesting that once a social hierarchy is established, individuals determined their sexual phenotype, regardless of initial sex, based on a simple operational principle: if subordinate express female, if dominant or not subordinate express male. This work demonstrates that similar mechanisms underlie sex change in both directions in L. dalli and potentially other sex changing species.     

Effects of 11-ketotestosterone on genital papilla morphology in the sex changing fish Lythrypnus dalli

Five days after implantation with 11-ketotestosterone (KT) female Lythrypnus dalli exhibited elevated urinary KT levels and male-like genital papilla morphology. Papilla length increased, width decreased, and length : width ratio increased. Control fish exhibited increases in papilla length, and length : width ratio on day 3, but not on day 5. Changes in the total area of the papilla were not statistically significant. It is suggested that KT mediates genital papilla morphology rapidly and thus may be a useful indicator of circulating KT in L. dalli.     

Horn growth rate and longevity: implications for natural and artificial selection in thinhorn sheep (Ovis dalli)

We used horn measurements from natural and hunted mortalities of male thinhorn sheep Ovis dalli from Yukon Territory, Canada, to examine the relationship between rapid growth early in life and longevity. We found that rapid growth was associated with reduced longevity for sheep aged 5 years and older for both the hunted and natural mortality data sets. The negative relationship between growth rate and longevity in hunted sheep can at least partially be explained by morphologically biased hunting regulations. The same trend was evident from natural mortalities from populations that were not hunted or underwent very limited hunting, suggesting a naturally imposed mortality cost directly or indirectly associated with rapid growth. Age and growth rate were both positively associated with horn size at death for both data sets, however of the two growth rate appeared to be a better predictor. Large horn size can be achieved both by individuals that grow horns rapidly and by those that have greater longevity, and the trade-off between growth rate and longevity could limit horn size evolution in this species. The similarity in the relationship between growth rate and longevity for hunted and natural mortalities suggests that horn growth rate should not respond to artificial selection. Our study highlights the need for the existence and study of protected populations to properly assess the impacts of selective harvesting.     

Muskox lungworm (Umingmakstrongylus pallikuukensis) does not establish in experimentally exposed thinhorn sheep (Ovis dalli)

Muskoxen (Ovibos moschlatus moschatus) on the northwestern mainland of Nunavut and Northwest Territories, Canada, are infected with the protostrongylid lungworm, Umingmaksrongylus pallikuuhkensis. The geographic range of this muskox population is expanding to the south and west, and it is anticipated that these animals will eventually become sympatric with Dall's sheep (Ovis dalli dalli) in the Mackenzie and Richardson Mountains. To address the concern of wildlife managers that U. pallikulkensis may infect and adversely affect Dalls sheep, four Dalls/Stone's (Ovis dalli stonei) hybrid lambs and one adult muskox (Ovibos moschatus wardi) were each given 100 third-stage larvae of U. pallikuukensis. All animals were intensively monitored for 9 mo postinfection (PI) using clinical examinations, fecal analyses, hematology, blood chiemistry and medical imaging. No first-stage larvae of U. pallikuinkensis were recovered from the lambs and monitoring revealed nio evidence that the parasite had established in any of these animals. First-stage larvae were found in the feces of the muskox beginning at 94 days PI, and typical parasite cysts were visible in lung radiographs at 188 days PI. This study addresses an important management and wildlife health issue associated with the potential for host-switching of pathogens and indicates that it is improbable that thinhorn sheep are suitable hosts for U. pallikuukensis.     

An integrative approach to sex change: social, behavioural and neurochemical changes in Lythrypnus dalli (Pisces)

This study examined three aspects of protogynous sex change in Lythrypnus dalli (Gobiidae): (1) social influences on the rate of sex change, (2) the sequence of behavioural changes, and (3) neuroendocrine changes. Social groups consisted of either four females, or four females with a male who was subsequently removed. Sex change occurred most rapidly in male- removed groups when the sex changer was larger than other females. Sex changers in female only groups and sex changers not larger than other females in male-removed groups changed sex at similar rates. These differences may be explained by two factors that affect dominance: prior knowledge of the social group and greater size. Sex changers were dominant to other females prior to male removal, and larger sex changers increased displacement rates three-fold immediately after male removal. Sex changers in the other groups did not show this increase in displacements. This early establishment of dominance accounts for the overall difference in the rate of sex change. Prior to spawning, however, all sex changers increased displacements and performed male-typical displays. Arginine vasotocin-immunoreactive forebrain cells of sex changers were similar in size to field-collected males, and larger than field-collected females. Previously nesting males also changed sex in male-only groups, but at slow rates. These data are combined with those of existing studies to generate an integrative model of sex change in this goby. Received: 17 March 1999 / Received in revised form: 15 May 1999 / Accepted: 28 May 1999     

Detecting the signature of selection on immune genes in highly structured populations of wild sheep (Ovis dalli)

The confounding effects of population structure complicate efforts to identify regions of the genome under the influence of selection in natural populations. Here we test for evidence of selection in three genes involved in vertebrate immune function - the major histocompatibility complex (MHC), interferon gamma (IFNG) and natural resistance associated macrophage polymorphism (NRAMP) - in highly structured populations of wild thinhorn sheep (Ovis dalli). We examined patterns of variation at microsatellite loci linked to these gene regions and at the DNA sequence level. Simple Watterson's tests indicated balancing selection at all three gene regions. However, evidence for selection was confounded by population structure, as the Watterson's test statistics from linked markers were not outside of the range of values from unlinked and presumably neutral microsatellites. The translated coding sequences of thinhorn IFNG and NRAMP are fixed and identical to those of domestic sheep (Ovis aries). In contrast, the thinhorn MHC DRB locus shows significant evidence of overdominance through both an excess of nonsynonymous substitution and trans-species polymorphism. The failure to detect balancing selection at microsatellite loci linked to the MHC is likely the result of recombination between the markers and expressed gene regions.