Identification by polymerase chain reaction (PCR) of Marshallagia marshalli and Ostertagia gruehneri from Svalbard reindeer

A polymerase chain reaction (PCR) assay to identify two common abomasal nematodes Marshallagia marshalli and Ostertagia gruehneri of Svalbard reindeer was developed. Species-specific PCR primers were designed from internal transcribed spacer (ITS)-2 sequences of rDNA and validated using morphologically identified adult male and female nematodes. Using the species-specific primers, a 110 bp fragment was amplified from M. marshalli and its minor morph Marshallagia occidentalis and a 149 bp fragment was amplified from Ostertagia gruehneri and its minor morph Ostertagia arctica. No PCR products were amplified from the third rare species, Teladorsagia circumcincta, or DNA from the reindeer host. The assay provides a useful tool to estimate species composition for both sexes in this nematode community.     

Obligate larval inhibition of Ostertagia gruehneri in Rangifer tarandus? Causes and consequences in an Arctic system

SUMMARY Larval inhibition is a common strategy of Trichostrongylidae nematodes that may increase survival of larvae during unfavourable periods and concentrate egg production when conditions are favourable for development and transmission. We investigated the propensity for larval inhibition in a population of Ostertagia gruehneri, the most common gastrointestinal Trichostrongylidae nematode of Rangifer tarandus. Initial experimental infections of 4 reindeer with O. gruehneri sourced from the Bathurst caribou herd in Arctic Canada suggested that the propensity for larval inhibition was 100%. In the summer of 2009 we infected 12 additional reindeer with the F1 and F2 generations of O. gruehneri sourced from the previously infected reindeer to further investigate the propensity of larval inhibition. The reindeer were divided into 2 groups and half were infected before the summer solstice (17 June) and half were infected after the solstice (16 July). Reindeer did not shed eggs until March 2010, i.e. 8 and 9 months post-infection. These results suggest obligate larval inhibition for at least 1 population of O. gruehneri, a phenomenon that has not been conclusively shown for any other trichostrongylid species. Obligate inhibition is likely to be an adaptation to both the Arctic environment and to a migratory host and may influence the ability of O. gruehneri to adapt to climate change.     

The population dynamics of Ostertagia gruehneri in reindeer: a model for the seasonal and intensity dependent variation in nematode fecundity

The gastrointestinal nematode Ostertagia gruehneri is a parasite of reindeer that can have a significant impact on host population dynamics. To gain a better understanding of the population dynamics of O. gruehneri, we parameterise a model for its fecundity that describes the observed seasonal and intensity dependent pattern of faecal egg counts well. The faecal egg count model is combined with a model for the seasonal faecal production rate of Svalbard reindeer to obtain quantitative estimates of the fecundity of O. gruehneri. The model is used to evaluate the relative contribution to pasture contamination of variation in the abundance of O. gruehneri and variation in reindeer densities. It is concluded that due to the intensity dependence in nematode fecundity, variation in reindeer population densities is likely to be the most important of these factors for pasture contamination.     

Development and availability of the free-living stages of Ostertagia gruehneri, an abomasal parasite of barrenground caribou (Rangifer tarandus groenlandicus), on the Canadian tundra

Climate change in the Arctic is anticipated to alter the ecology of northern ecosystems, including the transmission dynamics of many parasite species. One parasite of concern is Ostertagia gruehneri, an abomasal nematode of Rangifer ssp. that causes reduced food intake, weight loss, and decreased pregnancy rates in reindeer. We investigated the development, availability, and overwinter survival of the free-living stages of O. gruehneri on the tundra. Fecal plots containing O. gruehneri eggs were established in the Northwest Territories, Canada under natural and artificially warmed conditions and sampled throughout the growing season of 2008 and the spring of 2009. Infective L3 were present 3-4 weeks post-establishment from all trials under both treatments, except for the trial established 4 July 2008 under warmed conditions wherein the first L3 was recovered 7 weeks post-establishment. These plots were exposed to significantly more time above 30°C than the natural plots established on the same date, suggesting a maximum temperature threshold for development. There was high overwinter survival of L2 and L3 across treatments and overwintering L2 appeared to develop to L3 the following spring. The impact of climate change on O. gruehneri is expected to be dynamic throughout the year with extreme maximum temperatures negatively impacting development rates.     

Similarities between species of the Ostertagiinae (Nematoda: Trichostrongyloidea) in relation to host-specificity and climatic environment

Multivariate analyses of similarities between species of the Ostertagiinae were performed on the data from 585 publications. The environmental factors were 12 climates and host-specificity in ruminants (40 species) for 16 species of ostertagiines. This approach can be a useful tool for distinguishing morphs from species. The following pairs (species and its possible morph) of ostertagiines are proposed: Ostertagia ostertagi — O. lyrata, O. leptospicularis — O. kolchida, O. gruehneri — O. arctica, Teladorsagia circumcincta — T. trifurcata, Marshallagia marshalli — M. occidentalis, Spiculopteragia boehmi — S. mathevossiani, and S. asymmetrica — S. quadrispiculata.     

Resistance to abomasal nematodes and individual genetic variability in reindeer

Resistance to parasites is believed to have a widespread influence on demographic and adaptive processes. In systems where parasites impose a fitness cost on their host, heterozygotes may be selected because they are more resistant to parasites than homozygotes. Our objective was to assess the relationships between genomewide individual heterozygosity and abomasal nematode burdens in female Svalbard reindeer (Rangifer tarandus platyrhynchus) after the effects of host age, locality, season, and year had been accounted for. Samples were obtained from 306 female reindeer that were culled and genotyped at nine microsatellite loci. Reindeer in our study populations are mainly parasitized by the gastrointestinal nematodes Ostertagia gruehneri and Marshallagia marshalli. The infection intensity of each parasite differed between subpopulations, and among host age classes, seasons and years. We found no significant relationships between abomasal worm burdens, or lumen and mucosa larvae, of either O. gruehneri or M. marshalli and individual heterozygosity (or mean d(2)) alone or in interactions with host age, locality, and year. Although we analysed one of the largest data set available to date on gastrointestinal nematodes of a wild ruminant, we used a typical data set of nine genetic neutral markers that may have had low power to detect heterozygosity-fitness correlations. We conclude that the proportion of the variance in parasite resistance explained by individual heterozygosity for neutral genetic markers is low in Svalbard reindeer and in vertebrates in general, and we suggest that the candidate-gene approach might be more fruitful for further research on gene-fitness correlations.     

The role of parasites in the dynamics of a reindeer population

Even though theoretical models show that parasites may regulate host population densities, few empirical studies have given support to this hypothesis. We present experimental and observational evidence for a host-parasite interaction where the parasite has sufficient impact on host population dynamics for regulation to occur. During a six year study of the Svalbard reindeer and its parasitic gastrointestinal nematode Ostertagia gruehneri we found that anthelminthic treatment in April-May increased the probability of a reindeer having a calf in the next year, compared with untreated controls. However, treatment did not influence the over-winter survival of the reindeer. The annual variation in the degree to which parasites depressed fecundity was positively related to the abundance of O. gruehneri infection the previous October, which in turn was related to host density two years earlier. In addition to the treatment effect, there was a strong negative effect of winter precipitation on the probability of female reindeer having a calf. A simple matrix model was parameterized using estimates from our experimental and observational data. This model shows that the parasite-mediated effect on fecundity was sufficient to regulate reindeer densities around observed host densities.     

First sign in Italy of Nematodirus roscidus, Apteragia quadrispiculata and Ostertagia drozdzi, found in Dama dama

The authors refer the first report in Italy of Nematodirus roscidus Railliet, 1911, Apteragia quadrispiculata Jansen, 1958 and Ostertagia drozdzi Jancev, 1977 from Dama dama. In fallow deer they found also: Trichostrongylus axei, Trichostrongylus colubriformis, Trichostrongylus vitrinus, Spiculopteragia asymmetrica, Skrjabinagia arctica, Oesophagostomum venulosum, Trichuris ovis. Trichostrongylus axei, Trichostrongylus colubriformis, Trichostrongylus vitrinus and Skrjabinagia arctica were not found before in Dama dama in Italy.     

Antibodies to ruminant alpha-herpesviruses and pestiviruses in Norwegian cervids

A serologic survey revealed that Norwegian populations of free-ranging reindeer (Rangifer tarandus tarandus), roe deer (Capreolus capreolus), red deer (Cervus elaphus), and moose (Alces alces) have been exposed to alpha-herpesviruses and pestiviruses. A total of 3,796 serum samples collected during the period 1993-2000 were tested in a neutralization test for antibodies against bovine herpesvirus 1 (BHV-1) or cervid herpesvirus 2 (CerHV-2), and 3,897 samples were tested by a neutralization test and/or enzyme-linked immunosorbent assay for antibodies against bovine viral diarrhea virus (BVDV). Antibodies against alpha-herpesvirus were found in 28.5% of reindeer, 3.0% of roe deer, and 0.5% of red deer, while all moose samples were negative. In reindeer, the prevalence of seropositive animals increased with age and was higher in males than females. Antibodies against BVDV were detected in 12.3% of roe deer, 4.2% of reindeer, 2.0% of moose and 1.1% of red deer. The results indicate that both alpha-herpesvirus and pestivirus are endemic in reindeer and pestivirus is endemic in roe deer in Norway. The viruses may be specific cervid strains. Seropositive red deer and moose may have become exposed as a result of contact with other ruminant species.     

Comparative morphology of Ostertagia mossi and Ostertagia dikmansi (trichostrongylidae) from Odocoileus virginianus and comments on other Ostertagia spp. from the Cervidae

The synlophes of Ostertagia mossi Dikmans, 1931 and O. dikmansi Becklund & Walker, 1968 were found to be identical supporting recognition of these as the major and minor morphotypes, respectively, of a putative species pair within the Ostertagiinae. The cervical synlophe included three narrowly spaced, continuous, parallel ridges laterally and three parallel but more widely spaced ridges in the dorsal and ventral fields (Type II and Type A patterns, respectively). The disposition of the synlophe in conjunction with a long oesophageo-intestinal valve, and attributes of the genital cone and spicules are characteristics shared among other Ostertagia spp. from cervids in the Holarctic region that allow recognition of a species group morphologically distinct from the Ostertagiinae of domestic ruminants. The morphological similarity of O. mossi/O. dikmansi, O. leptospicularis Assadov, 1953/O. kolchida Popova, 1937 and O. gruehneri Skrjabin, 1929/O. arctica Mitzkewitsch, 1929 is substantial, but these species can be differentiated by small but constant differences in the synlophe and genital complex of males; females cannot be easily distinguished. The morphological similarity of this cryptic complex of species and their relatively restricted distribution among cervids suggests a coevolutionary history for hosts and parasites in the Holarctic region.     

Experimental studies on the interaction between infections of Ostertagia leptospicularis and other bovine Ostertagia species

Experimental infections of calves were carried out with either isolates of predominantly Ostertagia ostertagi, pure O. leptospicularis or a mixed isolate of equal numbers of both these species. The total worms established on day 21 for the mixed species from a total inoculum of 100 000 infective larvae, was 1.2 times greater than from 100 000 larvae of the O. ostertagi isolate and 3.3 times that of the pure O. leptospicularis isolate. The increased establishment in the mixed inoculum referred to both O. ostertagi and O. leptospicularis (days 17 and 21). These differences were both highly significant (P less than 0.01). The severity of the pathological changes was also greater in the mixed infections. It is suggested that these findings must be taken into account when control measures involving alternate grazing of sheep and cattle are being employed.     

Mitochondrial DNA and microsatellite DNA variation in domestic reindeer (Rangifer tarandus tarandus) and relationships with wild caribou (Rangifer tarandus granti, Rangifer tarandus groenlandicus, and Rangifer tarandus caribou)

Reindeer (Rangifer tarandus tarandus) in Alaska are semidomestic livestock descended from 1280 animals introduced from Siberia, Russia, approximately 100 years ago. Genetic variation at 18 microsatellite DNA loci and the cytochrome b gene of mitochondrial DNA (mtDNA) was quantified in reindeer from Alaska, Siberia (Russia), and Scandinavia and compared with wild North American caribou. Mean sequence divergence among 15 mtDNA haplotypes in reindeer was 0.007 substitutions per nucleotide site, and reindeer mtDNA is polyphyletic with caribou mtDNA. Microsatellite allele and mtDNA haplotype frequencies are similar between Alaskan and Russian reindeer and differentiated between these and Scandinavian reindeer. The frequencies of microsatellite alleles and mtDNA haplotypes are different in reindeer and wild caribou (Rangifer tarandus granti, Rangifer tarandus groenlandicus, and Rangifer tarandus caribou). Alaskan reindeer have maintained a genetic variation comparable to that in Russia and differentiated from that of wild caribou, >100 years after their introduction to Alaska.     

Taxonomic revision of the genus <Emphasis Type="Italic">Orloffia</Emphasis> (Nematoda: Ostertagiinae) based on an ITS-2 rDNA study

A comparative study of ITS-2 rDNA of the nematodes of the genera Orloffia, Ostertagia, Teladorsagia, and Trichostrongylus was performed. The obtained results allow us to consider Orloffia, Ostertagia, and Teladorsagia as closely related but distinct genera. The results of the study indicate that O. bisonis is the only valid species in the genus Orloffia, which has two morphologically distinct forms of males: O. bisonis (major morph) and O. kasakhstanica (minor morph). The percentage of similarity of ITS-2 sequences was 83.3–84.5% between Orloffia spp. and Ostertagia ostertagi, 84.9–86.1% between Orloffia spp. and Teladorsagia circumcincta, and 73.8–74.6% between Orloffia spp. and Trichostrongylus colubriformis.     

Dermacentor albipictus (Acari, Ixodidae) on captive reindeer and free-ranging woodland caribou

Infestations of winter ticks (Dermacentor albipictus) on two captive reindeer (Rangifer tarandus tarandus) are reported and may be associated with increased grooming and alopecia. Over 400,000 ticks were recovered from one reindeer. Few ticks (less than 25 ticks/animal) were found on three free-ranging woodland caribou (Rangifer tarandus caribou).     

Escaping parasitism in the selfish herd: age, size and density-dependent warble fly infestation in reindeer

It has been suggested that animals may escape attack from mobile parasites by aggregating in selfish herds. A selfish herd disperses the risk of being attacked among its members and the per individual risk of parasite infection should therefore decrease with increasing animal density through the encounter–dilution effect. Moreover, in a selfish herd, dominant and agile animals should occupy the best positions and thereby receive fewer attacks compared to lower ranked animals at the periphery. We tested these predictions on reindeer ( Rangifer tarandus tarandus ) parasitized by warble flies ( Hypoderma tarandi ). Warble flies oviposit their eggs on reindeer during summer and induce strong anti-parasitic behavioural responses in the herds. In this period, reindeer are sexually segregated; females and calves form large and dense herds while males are more solitary. After hatching, the warble fly larvae migrate under the skin of their host where they encyst. In the present study encysted larvae were counted on newly slaughtered hides of male calves and 1.5 year old males from 18 different reindeer herds in Finnmark, northern Norway with large contrasts in reindeer density. In reindeer, body mass is correlated with fitness and social status and we hypothesized that individual carcass mass reflected the animal's ability to occupy the best positions within the herd. Larval abundance was higher among the 1.5 year old males than among the calves. For calves we found in accordance with the selfish herd hypothesis a negative relationship between larval abundance and animal density and between larval abundance and body mass. These relationships were absent for the 1.5 year old males. We suggest that these differences were due to different grouping behaviour where calves and females, but not males, aggregated in selfish herds where they escaped parasitism.     

Arctic life adaptation--I. The function of reindeer hemoglobin

1. The functional properties of hemoglobin from the reindeer (Rangifer tarandus tarandus L.) are characterized as a function of pH, temperature and organic phosphate concentration. 2. Alongside overall similarities shared with most vertebrate hemoglobins, hemoglobin from the reindeer shows significant differences with respect to the effect of both organic phosphates and chloride anions. 3. The limited effect of temperature on oxygen binding (delta H = -4 kcal/mol O2) could be regarded as an interesting case of molecular adaptation to extreme environmental conditions.     

Sexual dimorphism in the pelvic bones of Svalbard reindeer, Rangifer tarandus platyrhynchus

The determination of sex of carcasses of reindeer and caribou (Rangifer tarandus) is complicated because both sexes normally carry antlers. Growth curves of males and females diverge sufficiently so that the length of long bones can often indicate the sex of a carcass if its age at death is known; results, however, are not always conclusive. Pronounced sexual dimorphism was found in the breadth of the ventro-medial border (rim) of the acetabulum of adult Svalbard reindeer. Measurement of the rim of the acetabulum thus provides a quick and conclusive means for determining the sex of carcasses of these animals. Similar results have been found in several other species of ungulates. Reasons for the dimorphism and for the non-normal distribution of the data for both sexes in reindeer are discussed.     

The primary structure of hemoglobin from reindeer (Rangifer tarandus tarandus) and its functional implications

The primary structures of alpha- and beta-chains of hemoglobin from reindeer (Rangifer tarandus tarandus) were determined. Comparison of the reindeer hemoglobin sequence with those of human and bovine hemoglobins showed 50 and 29 substitutions per alpha beta dimer, respectively. The influence of replacements on the modulation of hemoglobin oxygen affinity by heterothopic ligands and temperature, as well as their importance on the structure-function relationships in hemoglobin are discussed.