Tyzzer's disease in muskrats: re-examination of specimens of hemorrhagic disease collected by Paul Errington

Organisms typical of those seen in Tyzzer's disease of muskrats (Ondatra zibethica) were found in tissue sections from two muskrats found dead in Iowa in 1947 by Paul Errington. These findings support further the hypothesis that Errington's and Tyzzer's diseases of muskrats are a single entity.     

Habitat suitability in muskrats: a test of the food limitation hypothesis

Data on habitat selection, overwinter survival, and litter survival rate of muskrats indicate that fitness differs among habitats in marsh environments. We measured nine nutrient-dependent phenotypic variables of muskrats and one independent assessment of forage quality during the ice-free period (May-October) from 1992 to 1994. Data were used to test the hypothesis that differences in food resources limit the demographic performance of muskrats in marginal versus prime habitats (food limitation hypothesis, FLH). None of the null predictions relating habitat quality with reproductive parameters could be rejected statistically. Adult body mass and length were greater in prime habitats, but fat content did not differ, which suggested that the difference was associated with larger structural size of dominant animals in prime habitats. Growth rate of weaned juveniles was not different among habitats, which represented the strongest evidence for rejecting the FLH. Faecal crude protein content, which was used as an index of quality of food ingested, was marginally different between the two most suitable habitats, but the direction of the difference did not support our prediction. Overall, the temporal variation in faecal crude protein content was similar among habitats. We conclude that food is not a key factor limiting the demographic performance of muskrats among habitats during the ice-free season.     

Heat increment of feeding and its thermoregulatory benefit in the muskrat (Ondatra zibethicus)

The calorigenic effect of feeding and its potential benefit in defraying thermoregulatory costs and attenuating immersion hypothermia of adult muskrats were investigated. A single session of feeding on aquatic vegetation was sufficient to raise the metabolic rate of muskrats for a period of at least 5 h. The peak postprandial rate of oxygen consumption averaged 1.42 times the level established for fasted animals, and the heat increment of feeding accounted for about 40% of the metabolizable energy intake of muskrats. There was no evidence of a postprandial rise in oxygen consumption of muskrats that entered water at 18–19°C after feeding. In aquatic trials, average and minimum steady-state oxygen consumption rates of fed muskrats were similar to, or even lower than values recorded from fasted animals, implying substitution of heat increment of feeding for thermoregulatory heat production. Our data did not support the hypothesis that heat increment of feeding retards body cooling in water. Net body temperature decline in water was actually higher in fed animals than in fasted controls. However, since previously fed muskrats also entered water at an elevated body temperature, the final body temperature (at 30 min immersion) was similar in all groups. These findings suggest that metabolic heat generated incidental to preimmersion feeding could provide a thermoregulatory benefit to muskrats by reducing the need for active thermogenesis in water.     

Seasonal variation in body composition and morphology of adult muskrats in central Saskatchewan, Canada

We quantified seasonal variation in body composition and morphology of adult muskrats ( Ondatra zibethicus ) inhabiting freshwater marsh environments in central Saskatchewan, Canada. The study areas were characterized by long and cold winters extending over six months during which muskrats were restricted to foraging under ice. A total of 162 adult muskrats were collected during nine sampling periods across the year. The large accumulation of fat reserves (16% of body mass) during winter and the concurrent decline in protein mass suggested a reduced maintenance requirement associated with the presence of energy-rich food resources. Dietary fibre content increased significantly during mid-summer and was manifested by changes in gut morphology. Mobilization of fat reserves during summer months by both sexes reflected high energetic demands for reproduction. Males depleted fat reserves soon after spring break-up, while near-exhaustion of fat reserves in females occurred 4–6 weeks later, during lactation. Pregnant females contained significantly greater fat and protein reserves compared to non-pregnant and lactating females. The dynamics of body reserves in muskrats should be viewed as an integral part of the sex-specific life-history traits of this important herbivore species of marsh environments.     

Cross-species transmission of Giardia spp.: inoculation of beavers and muskrats with cysts of human, beaver, mouse, and muskrat origin

Giardia cysts isolated from humans, beavers, mice, and muskrats were tested in cross-species transmission experiments for their ability to infect either beavers or muskrats. Giardia cysts, derived from multiple symptomatic human donors and used for inoculation of beavers or muskrats, were shown to be viable by incorporation of fluorogenic dyes, excystation, and their ability to produce infections in the Mongolian gerbil model. Inoculation of beavers with 5 x 10(5) Giardia lamblia cysts resulted in the infection of 75% of the animals (n = 8), as judged by the presence of fecal cysts or intestinal trophozoites at necropsy. The mean prepatent period was 13.1 days. An infective dose experiment, using 5 x 10(1) to 5 x 10(5) viable G. lamblia cysts collected by fluorescence-activated cell sorting, demonstrated that doses of between, less than 50, and less than 500 viable cysts were required to produce infection in beavers. Scanning electron microscopy of beaver small intestine revealed that attachment of G. lamblia trophozoites produced lesions in the microvillous border. Inoculation of muskrats with G. lamblia cysts produced infections when the dose of cysts was equal to or greater than 1.25 x 10(5). The inoculation of beavers with Giardia ondatrae or Giardia muris cysts did not produce any infection; however, the administration to muskrats of Giardia cysts of beaver origin resulted in the infection of 62% of the animals (n = 8), with a prepatent period of 5 days. Our results demonstrated that beavers and muskrats could be infected with Giardia cysts derived from humans, but only by using large numbers of cysts.(ABSTRACT TRUNCATED AT 250 WORDS)     

Cross-species transmission of Giardia spp.: inoculation of beavers and muskrats with cysts of human, beaver, mouse, and muskrat origin.

Giardia cysts isolated from humans, beavers, mice, and muskrats were tested in cross-species transmission experiments for their ability to infect either beavers or muskrats. Giardia cysts, derived from multiple symptomatic human donors and used for inoculation of beavers or muskrats, were shown to be viable by incorporation of fluorogenic dyes, excystation, and their ability to produce infections in the Mongolian gerbil model. Inoculation of beavers with 5 x 10(5) Giardia lamblia cysts resulted in the infection of 75% of the animals (n = 8), as judged by the presence of fecal cysts or intestinal trophozoites at necropsy. The mean prepatent period was 13.1 days. An infective dose experiment, using 5 x 10(1) to 5 x 10(5) viable G. lamblia cysts collected by fluorescence-activated cell sorting, demonstrated that doses of between, less than 50, and less than 500 viable cysts were required to produce infection in beavers. Scanning electron microscopy of beaver small intestine revealed that attachment of G. lamblia trophozoites produced lesions in the microvillous border. Inoculation of muskrats with G. lamblia cysts produced infections when the dose of cysts was equal to or greater than 1.25 x 10(5). The inoculation of beavers with Giardia ondatrae or Giardia muris cysts did not produce any infection; however, the administration to muskrats of Giardia cysts of beaver origin resulted in the infection of 62% of the animals (n = 8), with a prepatent period of 5 days. Our results demonstrated that beavers and muskrats could be infected with Giardia cysts derived from humans, but only by using large numbers of cysts.(ABSTRACT TRUNCATED AT 250 WORDS)     

An outbreak of "wet tail" in hamsters due to Tyzzer's organism (author's transl)

In a colony of Golden hamsters, many animals were found to show signs of "wet tail", and spontaneous deaths occurred during two months. With this outbreak of wet tail and death, etiologic and pathologic studies were made on 11 cases supplied from the colony. Gross pathologic examination disclosed hemorrhagic enteritis in 10 and necrotic foci in the liver in 5 out of 10 cases examined. A number of bacilli morphologically suggestive of Tyzzer's organisms were detected on smears of cut surfaces of the affected intestines and livers. After inoculation with homogenate of the intestines from the moribund cases into hamsters by oral route, the same clinical manifestations as observed in spontaneous cases were reproduced, and the inoculated animals died in 10 to 14 days with numerous Tyzzer's organisms in their intestines and livers. When apparently healthy hamsters were introduced into contaminated cages with soiled bedding which were allowed to stand at room temperature for about 2 months, most of them developed signs of wet tail and died in 5 to 19 days having liver necrosis characteristic of Tyzzer's disease.     

Historical surveys reveal a long‐term decline in muskrat populations

The muskrat (Ondatra zibethicus) is an iconic species in Canada, valued for both its fur and its integral role in wetland ecosystems, and widely regarded for its perseverance. However, the resilience of this semiaquatic mammal seems to be in question now as increasing evidence points to widespread population declines. Recent analyses of harvest data across North America suggest a reduction in their numbers, but this has not been widely corroborated by population surveys. In this study we replicated historic muskrat house count surveys at two large Great Lakes coastal wetlands and present confirmation that declines in muskrat harvest correspond to actual declines in muskrat abundance. At the Point Pelee National Park marsh and the Matchedash Bay‐Gray Marsh wetland we found that mean muskrat house counts declined by 93% and 91% respectively between historic surveys 40–50 yrs ago and contemporary surveys over the past 7 yrs. The factors responsible for these dramatic declines remain unclear but there may be a relationship with changes in the habitat quality of these wetlands that have occurred over the same time frame. Not only is the loss of muskrats an issue for the resulting loss of the wetland ecosystem services they provide, but it may be an indication of broader marsh ecosystem degradation. As such, a scarcity of muskrats should be considered a red flag for the state of biodiversity in our wetlands. Continued surveys and ongoing research are needed to shed more light on the current status of muskrat populations and their marsh habitats across their native range.     

Prevalence of Giardia spp. in beaver and muskrat populations in northeastern states and Minnesota: detection of intestinal trophozoites at necropsy provides greater sensitivity than detection of cysts in fecal samples.

Surveys of the prevalence of the intestinal protozoan Giardia spp. in animal populations have relied almost exclusively on the detection of cysts in fecal samples. We have determined the prevalence of Giardia spp. in beaver and muskrat populations in four northeastern states and Minnesota by using both the detection of trophozoites in mucosal scrapings from live-trapped animals at necropsy and the detection of cysts in fecal samples collected from kill-trapped animals. In muskrats the prevalence of Giardia infection was 36.6% by cyst detection in fecal samples (n = 790) from kill-trapped animals and 95.9% in live-trapped muskrats when the intestinal contents were analyzed for the presence of trophozoites (n = 219). Similarly, in beavers, Giardia infection was 9.2% by cyst detection in fecal samples (n = 662) from kill-trapped beavers and 13.7% in live-trapped animals examined for the presence of intestinal trophozoites (n = 302). The detection of trophozoites in mucosal scrapings from live-trapped animals consistently yielded a significantly higher prevalence for both muskrats and beavers than did the method based on detection of cysts in the fecal samples. The prevalence of Giardia infection in juvenile and adult live-trapped muskrats was similar (92.5 and 94.4%, respectively), but the prevalence in juvenile live-trapped beavers (23.2%) was significantly greater than that seen in the adult animals (12.6%). No difference in Giardia prevalence on the basis of sex was seen in either animal species. Regional variation, often statistically significant, was seen in the prevalence of Giardia in beavers in the northeastern states and Minnesota, but was not detected for muskrats.     

The muskrat in biomedical research

Muskrats are aquatic rodents of moderate size which are plentiful throughout North America, but are not used commonly in the laboratory. Recently, we tested the feasibility of muskrats as experimental models and have found them to be acquired and cared for easily in conventional laboratory animal facilities. Some of their natural characteristics and diseases are described. The husbandry techniques that we used are presented and form a base for the preparation of future guidelines for the maintenance and use of feral animals in research. The results of some initial experiments testing the muskrat's utility for investigations of cardiorespiratory control mechanisms also are presented. Our data show that even anesthetized muskrats possess brisk and dramatic cardiovascular and respiratory reflexes. Our findings that their brains possess the cytoarchitectural and myeloarchitectural features comparable to other mammals, combined with their relative uniformity in size, has allowed us to locate specific neuronal loci stereotaxically. We suggest that the muskrat be considered as an experimental animal model for studies of the neural control of cardiorespiratory systems.     

Naturally occurring Bacillus piliformis infection (Tyzzer's disease) in guinea pigs

Two juvenile male Hartley guinea pigs (Cavia porcellus) were found dead 36 hours after receipt from a commercial source. Both guinea pigs had dependent, subcutaneous edema and excess serous fluid in their thoracic and abdominal cavities. Their livers were mottled and the cecal walls were reddened focally. Histopathologic exam revealed nests of Bacillus piliformis within the absorptive epithelial cells of the ileum, cecum and colon. Vegetative organisms and spore-like structures were observed in the cytoplasm of intestinal epithelial cells by electron microscopy. A diagnosis of Tyzzer's disease was made.     

Prevalence of Giardia spp. in beaver and muskrat populations in northeastern states and Minnesota: detection of intestinal trophozoites at necropsy provides greater sensitivity than detection of cysts in fecal samples

Surveys of the prevalence of the intestinal protozoan Giardia spp. in animal populations have relied almost exclusively on the detection of cysts in fecal samples. We have determined the prevalence of Giardia spp. in beaver and muskrat populations in four northeastern states and Minnesota by using both the detection of trophozoites in mucosal scrapings from live-trapped animals at necropsy and the detection of cysts in fecal samples collected from kill-trapped animals. In muskrats the prevalence of Giardia infection was 36.6% by cyst detection in fecal samples (n = 790) from kill-trapped animals and 95.9% in live-trapped muskrats when the intestinal contents were analyzed for the presence of trophozoites (n = 219). Similarly, in beavers, Giardia infection was 9.2% by cyst detection in fecal samples (n = 662) from kill-trapped beavers and 13.7% in live-trapped animals examined for the presence of intestinal trophozoites (n = 302). The detection of trophozoites in mucosal scrapings from live-trapped animals consistently yielded a significantly higher prevalence for both muskrats and beavers than did the method based on detection of cysts in the fecal samples. The prevalence of Giardia infection in juvenile and adult live-trapped muskrats was similar (92.5 and 94.4%, respectively), but the prevalence in juvenile live-trapped beavers (23.2%) was significantly greater than that seen in the adult animals (12.6%). No difference in Giardia prevalence on the basis of sex was seen in either animal species. Regional variation, often statistically significant, was seen in the prevalence of Giardia in beavers in the northeastern states and Minnesota, but was not detected for muskrats.     

Tyzzer's disease in free-living cottontail rabbits (Sylvilagus floridanus) in Maryland

Complement-fixing (CF) antibody to Bacillus piliformis antigen was found in 9 of 14 (64%) serum samples obtained from cottontail rabbits (Sylvilagus floridanus) killed in the wild. CF antibody was not present in the serum of 8 cottontail rabbits trapped as juveniles in the same geographic areas and held in captivity for 4 years. Sero-negative cottontail rabbits died acutely with lesions typical of Tyzzer's disease following the intragastric administration of 10(3.8) ELD50 of B. piliformis spores. The possible influence of Tyzzer's disease upon the cyclic population pattern of cottontail rabbits in the wild is discussed. A hypothesis is presented that B. piliformis spores passed in the feces of diseased wild animals could contaminate pastures, hay and grain, and thereby serve as sources of infection to other animals.     

Naturally occurring Tyzzer's disease in cotton-top tamarins (Saguinus oedipus)

We noted naturally occurring infection with Clostridium piliforme (Tyzzer's disease) in 2 captive-reared cotton-top tamarins (Saguinus oedipus). Spontaneous Tyzzer's disease has been reported in multiple species of laboratory, domestic, and wild animals but is extremely rare in humans and nonhuman primates. Distinct from idiopathic colitis, which is common in cotton-top tamarins, these 2 tamarins had severe, transmural, necrotizing typhlocolitis accompanied by myocarditis and hepatitis. Abundant bacteria compatible with C. piliforme, the etiologic agent of Tyzzer's disease, were present adjacent to lesions in the cecum-colon, liver, and heart. Therefore, colitis caused by C. piliforme, although rare, should be included as a differential diagnosis in cotton-top tamarins and as a cause of postnatal mortality in this species.     

Density-dependent habitat selection in muskrats: a test of the ideal free distribution model

Summary Two predictions of the ideal free distribution model, a null hypothesis of habitat selection, were examined using free-ranging muskrats. We rejected the prediction that the proportion of the animals found in each of five habitats was independent of population size. Data on over-winter occupancy of muskrat dwellings tend also to refute the prediction of equal fitness reward among habitats. Habitat type and water-level had a profound effect on the suitability of a site for settlement. We concluded that the observed pattern of muskrat distribution followed more closely an ideal despotic distribution where some individuals benefited from a higher fitness because of resource monopolization. Current theories of density-dependent habitat selection, which assume an ideal free distribution, would not apply to muskrats and possibly to many other mammal species.     

Enterohepatitis in Mongolian gerbils (Meriones unguiculatus) inoculated perorally with Tyzzer's organism (Bacillus piliformis)

Enterohepatitis was produced in Mongolian gerbils by intragastric inoculation with Tyzzer's organism from natural infection of a gerbil. Death occurred in 50 to 60% animals 5 to 7 days postinoculation (p.i.). On day 3 p.i., when a few necrotic foci appeared in the liver, a large amount of bacterial antigen was present within ileocecal enterocytes and reticuloendothelial cells of the Peyer's patches. Neutrophil and monocyte infiltration was found in the lamina propria. On day 5 or 6 p.i. there was severe necrotizing and hemorrhagic ileotyphlocolitis. Bacterial antigen was abundant within not only enterocytes, but also smooth muscle cells of the ileum and jejunum as well as reticular cells of the mesenteric lymph nodes. On day 7 p.i. the intestinal lesions subsided in the presence of fewer bacteria, while necrotizing hepatitis became well developed. The results indicated that Mongolian gerbils were highly susceptible to the oral route of infection with the Tyzzer's organism.     

Morbidity and mortality factors in key deer (Odocoileus virginianus clavium)

The population health of endangered Key deer (Odocoileus virginianus clavium) was monitored from 10 February 1986 to 28 September 2000 by necropsy of animals that were killed by vehicles, euthanized because of terminal injuries or disease conditions, or found dead. The predominant mortality factor during the period was collision with motor vehicles; however, several infectious diseases were diagnosed, including infections with Arcanobacterium pyogenes, Haemonchus contortus, Salmonella spp., and Mycobacterium avium subsp. paratuberculosis. During the period monitored, the only infectious disease that was thought to have affected population dynamics was haemonchosis. Nevertheless, several of the observed diseases have potential to impact viability of the Key deer population under appropriate environmental conditions.     

Detection of serum antibodies to Bacillus piliformis in mice and rats using an enzyme-linked immunosorbent assay.

Rats and mice were infected with Bacillus piliformis organisms at a dosage which resulted in clinical signs of Tyzzer's disease in gerbils. Although rats and mice did not show clinical signs of disease, rising antibody titers to B. piliformis were detected by enzyme-linked immunosorbent assay (ELISA) 2 to 6 weeks post-inoculation and remained at positive levels 11 weeks post-inoculation. Western blot analyses of sera from experimentally infected animals revealed banding patterns nearly identical to those obtained using hyperimmune serum. Results indicated that elevated ELISA titers reflected production of specific antibodies directed against antigens of B. piliformis. ELISA and Western blot analyses of naturally infected animals yielded similar results. These findings suggest that immunoassays such as ELISA can be used to detect subclinically infected rats and mice in the absence of clinical signs or histopathologic evidence of Tyzzer's disease.     

The roles of elevation and salinity as primary controls on living foraminiferal distributions: Cowpen Marsh, Tees Estuary, UK

Benthic foraminiferal assemblages in subrecent deposits are commonly used to reconstruct past sea level. Interpretations are generally made by comparison with either modern dead or total (live plus dead) assemblages. In both cases there will have been post-mortem changes that have differentially affected preservation. It is therefore important to establish the primary ecological controls by analysis of the living assemblages. We have determined the spatial and temporal variability of intertidal benthic foraminifera in the surface (0–1 cm) sediments from a time series survey of 31 sampling stations at Cowpen Marsh, for a period of 12 months. We counted 112,067 live foraminifera assigned to 28 species. The fauna was dominated by two agglutinated species (Jadammina macrescens and Trochammina inflata) on the high and middle marshes, and three calcareous species (Elphidium williamsoni, Haynesina germanica and Quinqueloculina spp.) on the low marsh and tidal flat.The standing crop of the whole intertidal zone, including the high, middle, low marsh and tidal flat habitats, and the individual species varied both temporally and spatially. The standing crop of the intertidal zone as a whole was greatest in the summer months and showed a positive correlation with elevation. The standing crops of the high and middle marshes showed similar temporal variation with peaks in summer and autumn and a trough in winter. The low marsh showed numerous peaks and troughs of standing crop during the year, whereas the tidal flat showed a single peak in summer. The standing crops of Jadammina macrescens and Trochammina inflata on the high and middle marshes peaked from April to May and August to October with troughs in winter. These agglutinated species showed a strong correlation with elevation. Haynesina germanica peaked in May to August and November to January on the low marsh, whereas on the tidal flat there was a single peak in July. The standing crops of E. williamsoni on the low marsh and tidal flat were relatively high in June and May, and July, respectively. Quinqueloculina spp. peaked in May to July on the low marsh and July on the tidal flat. The species was also found in the middle marsh from July to May and high marsh from September to November. Haynesina germanica showed a strong negative correlation with elevation, whereas the other two dominant calcareous species demonstrated weak negative correlations with both elevation and salinity.Reconstructing former sea level depends primarily on the recognition of high and middle marsh assemblages and in this study these are shown to be strongly controlled by elevation rather than salinity. Caution may be needed in interpreting low marsh and tidal flat data as salinity plays a more important role here.     

Body oxygen stores, aerobic dive limits, and the diving abilities of juvenile and adult muskrats (Ondatra zibethicus)

Intraspecific variability in body oxygen reserves, muscle buffering capacity, diving metabolic rate, and diving behavior were examined in recently captured juvenile and adult muskrats. Allometric scaling exponents for lung (b=1.04), blood (b=0.91), and total body oxygen storage capacity (b=1.09) did not differ from unity. The concentration of skeletal muscle myoglobin scaled positively with mass in 254-600-g juveniles (b=1.63) but was mass-independent in larger individuals. Scaling exponents for diving metabolic rate and calculated aerobic dive limit (ADL) were 0.74 and 0.37, respectively. Contrary to allometric predictions, we found no evidence that the diving abilities of muskrats increased with age or body size. Juveniles aged 1-2 mo exhibited similar dive times but dove more frequently than summer-caught adults. Average and cumulative dive times and dive&rcolon;surface ratios were highest for fall- and winter-caught muskrats. Total body oxygen reserves were greatest in winter, mainly due to an increase in blood oxygen storage capacity. The buffering capacity of the hind limb swimming muscles also was highest in winter-caught animals. Several behavioral indicators of dive performance, including average and maximum duration of voluntary dives, varied positively with blood hemoglobin and muscle myoglobin concentration of muskrats. However, none of the behavioral measures were strongly correlated with the total body oxygen reserves or ADLs derived for these same individuals.