Host-parasite associations of Eimeria spp. (Apicomplexa:Eimeriidae) in kangaroos and wallabies of the genus Macropus (Marsupialia:Macropodidae)

Faecal samples from 514 kangaroos and wallabies representing 12 species of the genus Macropus were examined for oocysts of Eimeria spp. Six species of Eimeria were redescribed from their type hosts, and on the basis of finding homologous oocysts in the faeces of other Macropus spp., host ranges for these coccidia were extended. Eimeria hestermani Mykytowycz, 1964 is redescribed from M. giganteus (eastern grey kangaroo) and is described from M. fuliginosus (western grey kangaroo), M. rufogriseus (red-necked wallaby), M. dorsalis (black-striped wallaby), and M. eugenii (tammar wallaby). E. toganmainensis Mykytowycz, 1964 is redescribed from M. rufus (red kangaroo) and the host range is extended to M. giganteus, M. fuliginosus, M. rufogriseus and M. eugenii. E. wilcanniensis Mykytowycz, 1964 is redescribed from M. rufus, and the host range is extended to M. giganteus, M. fuliginosus and M. robustus (euro or wallaroo). E. macropodis Wenyon & Scott, 1925 is redescribed from M. rufogriseus, and is described from M. giganteus, M. fuliginosus, M. rufus, M. irma (western brush wallaby), M. parryi (whip-tailed wallaby), M. dorsalis, M. eugenii, and M. parma (parma wallaby). E. fausti Yakimoff & Matschoulsky, 1936, E. cunnamullensis Mykytowycz, 1964 and E. purchasei Mykytowycz, 1964 are synonymized with E. macropodis. E. marsupialium Yakimoff & Matschoulsky, 1936 is redescribed from M. giganteus, and from M. fuliginosus. E. gungahlinensis Mykytowycz, 1964 is redescribed from M. fuliginosus, and from M. giganteus. Seven new species of Eimeria are described. E. flindersi, new species, is described from M. eugenii, M. rufogriseus, and M. antilopinus (antilopine wallaroo). E. prionotemni, new species, is described from M. eugenii, M. parryi, M. rufogriseus, M. agilis (agile wallaby) and M. dorsalis. E. mykytowyczi, new species, is described from M. agilis, M. antilopinus, and M. parryi. E. parryi, new species, is described from M. parryi. E. yathongensis, new species, is described from M. fuliginosus and M. giganteus. E. parma, new species, is described from M. parma, and E. desmaresti, new species, is described from M. rufogriseus. E. kogoni Mykytowycz, 1964, and E. rufusi Prasad, 1960 are considered species inquirendae. The host-parasite associations of these coccidia, and of similar species of Eimeria in other genera of Macropodoid marsupials, are discussed in relation to the postulated phylogeny of the hosts.     

Climatological correlates for body size of five species of Australian mammals

Size variation of body and skull of five species of Australian mammals (echidna, Tachyglossus aculeatus ; brush-tail possum, Trichosurus vulpecula ; eastern grey kangaroo, Macropus giganteus ; western grey kangaroo, M. fuliginosus ; red kangaroo, M. rufus ), is related to climatic factors. All five species show trends in body size that conform with Bergmann's rule, individuals from colder environments being larger than those from warmer areas. The western and eastern grey kangaroos also conform with Allen's rule, the relative size of their extremities being large in warmer areas. In four of the five species (not the red kangaroo) body size is also correlated with indices of biomass productivity. However, since biomass productivity and ambient temperature are related to some extent, it is difficult to separate the effects of these factors.     

Repertoires of social behaviour in captive and free-ranging grey kangaroos, Macropus giganteus and Macropus fuliginosus (Marsupialia: Macropodidae)

The eastern grey kangaroo, Macropus giganteus , and the western grey kangaroo, M. fuliginosus , are sibling species. The behaviour of 48 eastern grey kangaroos and 15 western grey kangaroos was observed for 249 h at Melbourne Zoo. Up to 31 eastern grey kangaroos and 38 western grey kangaroos were observed for 305 h where they occurred syntopically in the Grampians National Park.
In total, 46 social acts were identified in four social contexts between conspecifics. All but one of the acts were performed by both species in captivity, whereas fewer were seen performed by either species in the field. Five general acts were performed by all age/sex classes, and 10 acts occurred in mother-young contexts. High-intensity agonistic acts and all sexual acts were performed exclusively by adult males. In captivity, western grey males performed a truncated range of sexual acts directed at eastern grey females, but no interspecific sexual acts were performed by males of either species in the field, supporting the distinctiveness of the two species. The close evolutionary relationship between the two species was confirmed by high degree of concordance between their behavioural repertories, which differed only in the structure of the isolation call given by large pouch young.     

A nested PCR for the ssrRNA gene detects Trypanosoma binneyi in the platypus and Trypanosoma sp. in wombats and kangaroos in Australia

Trypanosome infections in their natural hosts are frequently difficult to detect by microscopy, and culture methods are unreliable and not suitable for all species of Trypanosoma. A nested PCR strategy for detecting and identifying Trypanosoma species, suitable for detecting both known and unknown trypanosomes, is presented. Thirty-two blood samples from 23 species of Australian birds and mammals were screened by a nested PCR for the presence of Trypanosoma sp. ssrRNA. Three infections were detected, one in an eastern grey kangaroo (Macropus giganteus), one in a common wombat (Vombatus ursinus) and one in a platypus (Ornithorhynchus anatinus). The kangaroo and wombat are new host records for Trypanosoma sp.; the platypus parasite was Trypanosoma hinneyi. The three parasites could be distinguished by restriction fragment length polymorphisms of the amplified fragment of the ssrRNA gene. The kangaroo and wombat parasites were also isolated in a semi-solid blood agar medium. The culture forms of the kangaroo trypanosome had an expanded flagellar sheath in which structures similar to hemidesmosomes were detected by EM. The nested PCR was at least as sensitive as culture, and analysis of the PCR products gave parasite-specific fingerprints. Therefore this method could be suitable for rapidly screening host animals for the presence of trypanosomes and identifying the infecting strain.     

Free-ranging heart rate, body temperature and energy metabolism in eastern grey kangaroos (Macropus giganteus) and red kangaroos (Macropus rufus) in the arid regions of South East Australia

Eastern grey kangaroos (Macropus giganteus) are generally regarded as mesic inhabitants. Even though access to drinking water in permanent stock watering troughs is commonly available, these animals are still found in only low densities in arid pastoral areas. We hypothesised that the differential success of red and grey kangaroos in the arid zone may be due to higher energy requirements of M. giganteus with a concomitant need for increased food, rather than limitations imposed by inadequate water access. We set out to test this by indirectly measuring energy expenditure through the monitoring of heart rate by radio telemetry in semi-free-ranging eastern grey and red kangaroos (Macropus rufus). Radio telemetry measurements of heart rate were calibrated against oxygen consumption and were used in the assessment of energy expenditure of animals maintained in an 8-ha enclosure in the arid zone of southeast Australia. Heart rate provided a reliable estimate of oxygen consumption. This well-correlated relationship was curvilinear and was established for each individual. Behavioural observations revealed that both kangaroo species spent most of the day in low energy demanding activities. M. rufus were more active at night whilst M. giganteus were more active in the early mornings and late afternoons. Like other marsupials, both species had low field metabolic rates (FMRs). However, M. giganteus in keeping with their mesic history had higher FMRs than the more arid-adapted M. rufus, particularly during water restriction. Body temperature telemeters revealed a further species difference in that under hot conditions when water is freely available, M. rufus exhibits a higher and more labile daytime body temperature than M. giganteus. During the hottest part of the day M. giganteus maintain body temperature, relying upon increased evaporative cooling mechanisms, such as licking. Indeed, only when access to drinking water was restricted was thermolability evident in M. giganteus. Differences in behaviour and concomitant energy expenditure may thus contribute substantially to the divergent distribution and abundance of these two kangaroo species.     

"Whey" proteins of milk of the red (Macropus rufus) and eastern grey (Macropus giganteus) kangaroo

1. A comparison is made of gel electrophoretic patterns of the "whey" proteins of the milk of red (Macropus rufus) and eastern grey (Macropus giganteus) kangaroos at various stages of lactation. Qualitative and quantitative changes occur with time during the mature phase of lactation of both types. Their onset is related solely to the stage of lactation. "Whey" proteins are isolated and characterised and the nature of protein changes determined for the first time. 2. The anodic electrophoretic pattern is divided into 6 main zones (designated A F in order of decreasing mobility) and 2 cathodic zones (G and H) that are only detected in the milk of M. giganteus. 3. Zones A, B and C are milk specific. Zone B is present throughout lactation in both species and is an alpha-lactalbumin. Zones A and C are present only in late lactation, zone C, usually, but not always, appearing first. Zone A is an alpha-lactalbumin in M. giganteus, but is not an alpha-lactalbumin in M. rufus. Zone C appears to be the same protein in both species and is possibly a beta-lactoglobulin. 4. Zone D is kangaroo serum albumin and zone E is possibly a beta 2-microglobulin. Zone F contains three main iron (III) binding bands whose relative intensity varies with stage of lactation. Their intensity differs from the corresponding blood serum transferrin bands. 5. Zone H of Macropus giganteus is a lysozyme. 6. Lactose is present in the milk, but is not the principal sugar. 7. The significance of the results is discussed.     

Ventilatory accommodation of oxygen demand and respiratory water loss in kangaroos from mesic and arid environments, the eastern grey kangaroo (Macropus giganteus) and the red kangaroo (Macropus rufus)

We studied ventilation in kangaroos from mesic and arid environments, the eastern grey kangaroo (Macropus giganteus) and the red kangaroo (Macropus rufus), respectively, within the range of ambient temperatures (T(a)) from -5 degrees to 45 degrees C. At thermoneutral temperatures (Ta=25 degrees C), there were no differences between the species in respiratory frequency, tidal volume, total ventilation, or oxygen extraction. The ventilatory patterns of the kangaroos were markedly different from those predicted from the allometric equation derived for placentals. The kangaroos had low respiratory frequencies and higher tidal volumes, even when adjustment was made for their lower basal metabolism. At Ta>25 degrees C, ventilation was increased in the kangaroos to facilitate respiratory water loss, with percent oxygen extraction being markedly lowered. Ventilation was via the nares; the mouth was closed. Differences in ventilation between the two species occurred at higher temperatures, and at 45 degrees C were associated with differences in respiratory evaporative heat loss, with that of M. giganteus being higher. Panting in kangaroos occurred as a graded increase in respiratory frequency, during which tidal volume was lowered. When panting, the desert red kangaroo had larger tidal volumes and lower respiratory frequencies at equivalent T(a) than the eastern grey kangaroo, which generally inhabits mesic forests. The inference made from this pattern is that the red kangaroo has the potential to increase respiratory evaporative heat loss to a greater level.     

Apparent lack of genetic variation within Pelecitus roemeri (Nematoda: Filarioidea) from three Australian species of macropodid marsupial.

An electrophoretic study of Pelecitus roemeri from Macropus robustus, M. giganteus and Wallabia bicolor revealed no genetic differences at 23 enzyme loci. The genetic data support the existing morphological evidence that P. roemeri from these three hosts represents a single species. The data show no genetic variation between nematodes from the same or different host species collected in northern and southern Australia. This result is discussed briefly in relation to Price's model of parasite speciation.     

Bacteriophages from the forestomachs of Australian marsupials.

Bacteriophages were observed in forestomach contents from three species of Australian macropodoid marsupials possessing a foregut fermentative digestion: the eastern grey kangaroo (Macropus giganteus), the eastern wallaroo (Macropus robustus robustus), and the rufous bettong (Aepyprymnus rufescens). Forty-six morphologically distinct phage types, representing the families Myoviridae, Siphoviridae, and Podoviridae, were identified. The range of forms varied between host species. The greatest diversity of phage types was found in forestomach contents of the wallaroo, and few phage types were recorded from the rufous bettongs. It is concluded that macropodoid marsupials, in common with their eutherian counterparts, possess diverse populations of bacteriophages in their fermentative forestomachs.     

Bacteriophages from the forestomachs of Australian marsupials.

Bacteriophages were observed in forestomach contents from three species of Australian macropodoid marsupials possessing a foregut fermentative digestion: the eastern grey kangaroo (Macropus giganteus), the eastern wallaroo (Macropus robustus robustus), and the rufous bettong (Aepyprymnus rufescens). Forty-six morphologically distinct phage types, representing the families Myoviridae, Siphoviridae, and Podoviridae, were identified. The range of forms varied between host species. The greatest diversity of phage types was found in forestomach contents of the wallaroo, and few phage types were recorded from the rufous bettongs. It is concluded that macropodoid marsupials, in common with their eutherian counterparts, possess diverse populations of bacteriophages in their fermentative forestomachs.     

Detection by allozyme electrophoresis of cryptic species of Hypodontus macropi (Nematoda: Strongyloidea) from macropodid marsupials.

Allozyme electrophoresis of 98 Hypodontus macropi from eight different species of hosts using 24 enzymes revealed a complex of at least six sibling species, with 15-50% fixed genetic differences between taxa. Except for the taxon parasitizing Macropus rufus/M. robustus, pairs of parasite taxa were, in each case, sympatric at each locality examined, thus supporting the conclusion that they represent valid species. The existence of a series of host-specific nematode taxa explains many of the inconsistencies noted previously in the host distribution of H. macropi. Comparison of parasite allozyme phenograms with host phylogeny suggests that four of the speciation events could be attributable to cospeciation and two to host switching. A clear case of host switching between M. rufus/M. robustus and M. fuliginosus was found.     

Intraspecific variation,sex-biased dispersal and phylogeography of the eastern grey kangaroo (Macropus giganteus)

Genetic information has played an important role in the development of management units by focusing attention on the evolutionary properties and genetics of populations. Wildlife authorities cannot hope to manage species effectively without knowledge of geographical boundaries and demic structure. The present investigation provides an analysis of mitochondrial DNA and microsatellite data, which is used to infer both historical and contemporary patterns of population structuring and dispersal in the eastern grey kangaroo (Macropus giganteus) in Australia. The average level of genetic variation across sample locations was one of the highest observed for marsupials (h=0.95, HE=0.82). Contrary to ecological studies, both genic and genotypic analyses reveal weak genetic structure of populations, where high levels of dispersal may be inferred up to 230 km. The movement of individuals was predominantly male-biased (average Nem=22.61, average Nfm=2.73). However, neither sex showed significant isolation by distance. On a continental scale, there was strong genetic differentiation and phylogeographic distinction between southern (TAS, VIC and NSW) and northern (QLD) populations, indicating a current and/or historical restriction of gene flow. In addition, it is evident that northern populations are historically more recent, and were derived from a small number of southern founders. Phylogenetic comparisons between M. g. giganteus and M. g. tasmaniensis indicated that the current taxonomic status of these subspecies should be revised as there was a lack of genetic differentiation between the populations sampled.     

A New Family of Entodiniomorph Protozoa from the Marsupial Forestomach, with Descriptions of a New Genus and Five New Species

ABSTRACT. A unique group of entodiniomorph protozoa was found in forestomach contents from quokka ( Setonix brachyurus ), western grey kangaroo ( Macropus fuliginosus ), red kangaroo ( Macropus rufus ) and euro ( Macropus robustus erubescens ). A new genus, Macropodinium n.g., containing five new species, is described. Three species are described from forestomach contents of the quokka: Macropodinium baldense n. sp., Macropodinium moiri n. sp. and Macropodinium setonixum n. sp. A single species, Macropodinium ennuensis n. sp., is described from the red kangaroo and euro. The last species, Macropodinium yalanbense n. sp., is described in forestomach contents from the western grey kangaroo. At least three distinct features in the new genus are incompatible with any of the described families in the order Entodiniomorphida. On this basis, the new family Macropodiniidae has been created.     

Molecular characterization of Eimeria species in macropods

A total of 597 faecal samples were collected from western grey kangaroos (Macropus fuliginosus), Euros (M. robustus), red kangaroos (M. rufus) in Western Australia and Eastern Grey Kangaroos (M. giganteus) from Victoria and screened for the presence of Eimeria by PCR at the 18S ribosomal RNA (rRNA) locus. The overall prevalence was 24.3% (145/597). At the 18S rRNA locus, sequences were obtained for 25 of the 145 positives. Phylogenetic analysis indicated that all the macropod-derived Eimeria species grouped in a separate marsupial clade that included Eimeria trichosuri from brushtail possums. At least 6 different clades were identified within the marsupial isolates and many of the genotypes identified are likely to be valid species, however morphological and biological data need to be collected to match sequences to previously characterized Eimeria species or identify if they are new species.     

The phenomenon of phylogenetic synhospitality in acariform mites (acari: acariformes)--the permanent parasites of vertebrates

The term synhospitality means the association of two or more closely related parasite species with one host species (Eichler, 1966). The cases of two or three synhospitalic species are known from the same host species, and especially ones where parasites were recorded from different parts of the host range, are quite common. The most ordinary reason causing synhospitality in permanent parasites is the host switching. Nevertheless, there are a number of synhospitality cases, where the parasite complex is monophyletic because evolved on a single host species. The special term--"phylogenetic synhospitality" (FS) is proposed for these cases of synhospitality. Most known cases of FS in acariform mites, permanent parasites of vertebrates, are analysed. It is found out that both astigmatan and prostigmatan parasite mites demonstrate a numbers of FS. The majority of these examples represent parasitism of two or three synhospitalic parasite species. Impressive examples of FS involving a number of synhospitalic species is shown by only astigmatan mites inhabiting the fur of mammals or plumage of birds. Most known examples involving four or more mite species are discussed: 51 mite species of the genus Schizocarpus (Chirodiscidae) parasitizing Castor fiber and C. canadensis (Castoridae); 6 species of Listrophorus spp. (Listrophoridae) from Ondatra zibethicus (Cricetidae); 23 species of Listrophoroides s. 1. (Atopomelidae) from Maxomys surifer (Muridae); 21 species of Cytostethum (Atomelidae) from Potorous tridactylus (Potoridae); 4 species of Listrophoroides (Afrolistrophoroides) from Malacomys longipes (Muridae); 7 species of Fainalges (Xolalgidae) from Aratinga holochlora (Psittacidae); 4 species of Zygepigynia (Pteronyssidae) from Chrysocolaptes lucidus (Picidae). The main reason of FS is that, in spite of the Fahrenholz's rule, the speciation of many parasites proceeds much more intensively than in their hosts because of the more rapid replacement of the parasitic generations. The first factor causing FS is the mite speciation it temporary segregated populations of the host (allopatric speciation). In this case, the "multispecies complexes" appeared after the subsequent reintegration of the host populations formerly isolated. The second factor is the speciation due to the specialization of mites to local microhabitats in the fur or plumage of host (sympatric or synxenic speciation). The second way of speciation is most characteristic for mites with highly specialized attaching structures. The phenomenon of FS more resides in ectoparasites of mammals rather than in feather mites in spite of much more structural complicacy of plumage rather than the fur. The high mobility of birds and wide dispersion of their new generations probably embarrass the process of sympatric speciation in their parasites. As a rule, only really significant geographical barriers play role for population isolation in birds. Thus, it could be concluded that two independent factors or their combination lead to FS. (i) The complex and/or disjunctive host range giving a possibility for allopatric speciation in parasites. (ii) The deep mite specialization to local microhabitats on the host body causing sympatric (synxenic) speciation. Fur of mammals and plumage of birds are very complicated in structure and microconditions and provide a considerable number of different microhabitats for mites inhabiting them. The prevalence of one of these two factors depends on the biological peculiarities of both parasites and their hosts. In mites with lesser specialized attaching organs, for example in atopomelids, allopatric speciation dominates. In mites with strongly specialized attaching organs, for example in listrophorids or chirodiscids, both pathways of speciation may take place. In feather mites, sympatric speciation should be more probable due to quite complicate and various structure of feathers in avian hosts. In fur mites, sympatric speciation is more likely in mites parasitizing hosts with peculiar ecology, for example in semiaquatic rodents possessing quite different fur structure in different parts of the body.     

Coevolution is linked with phenotypic diversification but not speciation in avian brood parasites

Coevolution is often invoked as an engine of biological diversity. Avian brood parasites and their hosts provide one of the best-known examples of coevolution. Brood parasites lay their eggs in the nests of other species, selecting for host defences and reciprocal counteradaptations in parasites. In theory, this arms race should promote increased rates of speciation and phenotypic evolution. Here, we use recently developed methods to test whether the three largest avian brood parasitic lineages show changes in rates of phenotypic diversity and speciation relative to non-parasitic lineages. Our results challenge the accepted paradigm, and show that there is little consistent evidence that lineages of brood parasites have higher speciation or extinction rates than non-parasitic species. However, we provide the first evidence that the evolution of brood parasitic behaviour may affect rates of evolution in morphological traits associated with parasitism. Specifically, egg size and the colour and pattern of plumage have evolved up to nine times faster in parasitic than in non-parasitic cuckoos. Moreover, cuckoo clades of parasitic species that are sympatric (and share similar host genera) exhibit higher rates of phenotypic evolution. This supports the idea that competition for hosts may be linked to the high phenotypic diversity found in parasitic cuckoos.     

A test of host-associated differentiation across the 'parasite continuum' in the tri-trophic interaction among yuccas, bogus yucca moths, and parasitoids

Parasitic taxa span an antagonistic continuum, with some parasites inflicting no fitness costs to some that kill the host after feeding. Host-associated differentiation is postulated as a major process facilitating speciation in many parasitic taxa. Here, I examined the importance of host-associated differentiation in a parasitoid wasp that develops on yucca moths in the genus Prodoxus. Prodoxus are specialists on Yucca , and moth speciation is closely tied to differences in microhabitat use within a plant and among host plant species. Parasitoids in the genus Eusandalum have been reared from Prodoxus species distributed across Yucca . Estimates of host-use patterns obtained through rearings of adult wasps were combined with surveys of mitochondrial DNA cytochrome oxidase I sequence data and amplified fragment length polymorphism markers to determine if populations of Eusandalum were genetically structured based on host use. Eusandalum populations were genetically structured based on geographical distance rather than moth host species, microhabitats within plants, or Yucca species. The results are contrary to the patterns observed in the host genus Prodoxus . Although parasitoids exhibit parasite-like characteristics, these results suggest that Eusandalum may be best viewed as a predator. Female wasps are able to utilize any moth species present at a given locality, and there is little likelihood that host specialization may facilitate population subdivision and speciation.     

Mortalities associated with herpesvirus infection in captive macropods

Fatal cases of herpesvirus infection in a quokka (Setonix brachyurus), eight grey dorcopsis wallabies (Dorcopsis muelleri luctuosa) in two separate outbreaks, as well as presumptive fatal herpesvirus infection in a western grey kangaroo (Macropus fuliginosus) are described. All were captive animals and deaths were preceded by a period of stress. Lesions occurred most often in alimentary tract epithelium, respiratory tract, skin, genital epithelium, conjunctivae, liver and adrenal cortex. Herpesviruses were recovered from the quokka and a grey dorcopsis wallaby from each outbreaks. The implications, particularly concerning certain apparently susceptible species, for those responsible for the health of macropod collections are discussed.     

Satellite species in lampreys: a worldwide trend for ecological speciation in sympatry?

Amongst several theories of speciation, sympatric speciation has been the most controversial but it is now widely accepted that populations can become reproductively isolated without being separated geographically. One problem with the acceptance of the theory of sympatric speciation, however, has been the lack of supporting empirical data and it is still believed that geographical isolation is responsible for the majority of speciation events. Here the example of species pairs in lampreys suggests that sympatric speciation in a whole taxonomic group could occur throughout its worldwide range. Lampreys occur in two ecologically distinct forms: parasitic mostly anadromous species that forage on tissue and body fluids of host fishes, and non‐parasitic forms that, apart from a short adult life when they cease feeding, spend their entire life as filter feeders in the substratum of stream beds. Both forms occur in sympatric species pairs throughout the range of lampreys that occur in Eurasia, North America and Australia and it is widely acknowledged that non‐parasitic forms derive from parasitic forms. The larvae of both forms can be distinguished by their potential fecundity and therefore, it is argued that the mode of life is not a consequence of different ecological conditions. Furthermore, as lampreys prefer to choose mates of similar sizes and fertilization success decreases with increasing difference in body size, there is a strong disruptive selection between the two forms and they are therefore reproductively isolated. Besides theoretical aspects, the similarity of the species pairs, together with their occurrence in sympatry, the occurrence of forms with intermediate characteristics, and examples where speciation might be in progress, hints at the possibility that speciation also occurred in sympatry. The difference between lampreys and other examples of sympatric speciation is that there seems to be a trend towards sympatric speciation events throughout the worldwide range of lampreys which is neither restricted to relatively small localities nor caused by human disturbance. Species pairs in lampreys therefore offer a unique possibility of studying the process of sympatric speciation on a large scale.