Aggressive interactions between three species of freshwater crayfish of the genus Cherax (Decapoda: Parastacidae)

The freshwater crayfish, Cherax destructor Clark, native to southeastern Australia, was first introduced to farm dams in southwestern Western Australia in 1932. The geographic range of the crayfish in Western Australia has increased substantially since then, and in recent years it has become established in natural waterways where it co-occurs with species of freshwater crayfish endemic to southwestern Australia, Cherax cainii and Cherax quinquecarinatus. The potential for competitive exclusion of these endemic species by C. destructor was investigated through laboratory experiments measuring aggressive behaviour. Body mass and species were found to be important factors governing aggressive dominance between C. cainii and C. destructor, with C. cainii winning significantly more interactions only when they were larger in body mass than their opponent. In trials between C. quinquecarinatus and C. destructor of similar body mass, there was no difference between the number of interactions 'won' by the two species. The implications for natural populations are discussed.     

Host effects on glutathione S-transferase activity in Fasciola hepatica

Glutathione S-transferases (GST, E.G. 2.5.1.18) in Fasciola hepatica from sheep were previously found to be extremely variable with regard to specific GST activity and isoenzyme profile within and between parasite isolates. The effect of the host on GST activity and isoenzyme profile was examined by infecting mice, rats and cattle as well as sheep with one or the other of two isolates—either salicylanilide-resistant or salicylanilide-susceptible F. hepatica. In the case of both isolates, GST activity in hosts relatively resistant to reinfection—rats and cattle—was lower and more restricted in range compared with hosts susceptible to multiple infection—mice and sheep. In the case of the rat flukes, there was little variation in isozyme profiles whereas cattle flukes appeared to exhibit more variation than sheep flukes. In mice, despite the apparent variability in GST activity, only one GST band was found in the isoenzyme profiles. Therefore, the host appears to exert a pronounced effect on the activity and expression of GSTs in F. hepatica which may be related to variation in the immune responses of the different hosts during infection.     

Experimental inoculation of meadow voles (Microtus pennsylvanicus), house mice (Mus musculus), and Norway rats (Rattus norvegicus) with Mycobacterium bovis

Mycobacterium bovis has a wide host range that includes several wildlife species, and this can hamper attempts to eradicate bovine tuberculosis from livestock. The purpose of this study was to determine if common rodent species, namely meadow voles (Microtus pennsylvanicus), house mice (Mus musculus), and Norway rats (Rattus norvegicus), that inhabit the bovine tuberculosis endemic area of Michigan, can be experimentally infected with M. bovis. The objectives of the study were: 1) to determine if these rodent species can be infected, and if so, to document attendant pathologic processes/pathogenesis; 2) to detect any fecal shedding of M. bovis; and 3) to evaluate the relative susceptibility of the three species to M. bovis infection. For each species (n=36) there were two treatment (n=12/group) and one or two control groups depending on species (n=6-12/group); the maximum study duration was 60 days. The meadow vole treatments consisted of high dose inocula that were given by oral or intranasal routes, whereas the house mice and Norway rats were given only oral inocula at either a high or low dose. Of the three species, meadow voles were most susceptible to M. bovis infection. Upon intranasal inoculation, all 12 voles were infected as determined by gross and microscopic lesions and culture of M. bovis from tissue and feces. Seven of the 12 meadow voles inoculated orally were infected. House mice also were susceptible; M. bovis was isolated from 14 of 24 animals. Only one Norway rat in the high dose treatment group was positive by culture and this was the only animal from which minimal attendant lesions were observed. Results of this study indicate that meadow voles and house mice can be infected with M. bovis and might serve as spillover hosts. Concerted efforts should, therefore, be made to reduce or eliminate these rodents on premises where M. bovis-infected livestock are present.     

Preliminary host specificity testing of Endophyllum osteospermi (Uredinales, Pucciniaceae), a biological control agent against Chrysanthemoides monilifera ssp. monilifera

Chrysanthemoides monilifera ssp. monilifera, indigenous to the Western Cape Province of South Africa, is a serious invader of native vegetation in south-eastern Australia. The rust fungus Endophyllum osteospermi causes witches' brooms on C. monilifera ssp. monilifera in South Africa, and is associated with a reduction in growth and seed production of its host under natural conditions, as well as mortality of severely infected bushes. This rust fungus is considered to be a potential biological control agent for use against C. monilifera ssp. monilifera in Australia. Endophyllum osteospermi has a long latent period, typically between 6 and 24 months between infection and the initiation of witches' brooms. This long latent period makes the logistics of doing traditional host specificity testing, in which all test plant species are inoculated and observed for symptom development, unfeasible for this rust fungus. Germination of aecidioid teliospores and penetration by basidiospores were observed on the surface of excised leaves of 32 test plant species at 4 days after inoculation, and compared to that on C. monilifera ssp. monilifera. Germinating aecidioid teliospores aborted on 14 test plant species, whilst no penetration was attempted on a further 12 test plant species. Penetration only occurred on nine of the 32 test plant species, in addition to C. monilifera ssp. monilifera. Inoculating whole plants of nine selected test plant species confirmed the above results. Therefore, only the test plant species in which penetration occurred, or at least was attempted, need to undergo comprehensive host specificity testing. Pending these results, E. osteospermi may be suitable for release in Australia for the biological control of C. monilifera ssp. monilifera.     

五种小哺乳动物活动节律的初步研究

在一昼夜中,各种动物的活动均有一定的周期性,其活动期和安静期常有规律地交替变换着。本文研究了五种小哺乳动物:中华鼢鼠（Myosalax fontanierii）高原鼠兔（Ochotona curzoniae）根田鼠（Microtus oeconomus）小家鼠（Mus musculus）和褐家鼠（Rattus norvegicus）的活动节律,前3种所见报道甚少。     

Factors influencing the relative abundance of invasive predators and omnivores on islands

Invasive alien species (IAS) are the major cause of native species extinctions on islands worldwide. To mitigate or eliminate IAS impacts, eradication is often the best alternative. However, IAS removal may result in cascading effects, through increase in prey abundance, mesopredator release, or competitor release. Our objective is to determine which ecological processes may influence the relative abundance of invasive carnivores and rodents on an insular system. We find that feral cat and mustelids relative abundance was strongly related by prey abundance, and for the feral cat, abundance was also controlled by habitat; these results suggest that bottom-up control through environmental filtering could be the mechanism explaining predator abundance. For rodents, we find that the abundance of the black rat was mostly controlled by the abundance of Norway rat and house mice, and food availability; the Norway rat by the abundance of black rat, a house mice and of mustelid predators; and house mouse by the other rodents and food availability. These results suggest that several mechanisms could be concurrently controlling abundance of these species; competition and predation for Norway rat, and competition and bottom-up control by environmental filtering for the other two rodents. While different factors explain the abundance of invasive species within the same functional group, food resource availability is, in general, the main controller of abundance of invasive rodents and carnivores in the Azores. Therefore, IAS management actions in these islands should focus on limiting the access to food resources and shelter, mainly near to human populations.

     

Multiple Cryptosporidium genotypes detected in wild black rats (Rattus rattus) from northern Australia

As part of a broader investigation into the potential role of black rats (Rattus rattus) as disease vectors into native small mammal populations of northern Australia, blood and faecal samples from wild black rats were screened by molecular methods, for piroplasms (Babesia and Theileria), trypanosomes and the enteric parasite Cryptosporidium. While piroplasms and trypanosomes were not detected in the blood of these animals, the overall prevalence of Cryptosporidium 18S rDNA in faecal samples was 8.2% (7/85). Co-occurrence of multiple genotypes was observed in 57.1% of the infected individuals (4/7); cloning and re-sequencing resulted in 14 sequences which broadly grouped with Cryptosporidium sp. rat-genotypes II and III. A novel rat-derived Cryptosporidium sp. genotype at the actin locus was also obtained from five animals. The relatively low infection rate detected, and the epidemiological data on cryptosporidiosis, do not conclusively support a current threat to native Australian mammals from black rats carrying Cryptosporidium. However, this observation is based on sampling limited isolates, in limited regions. Further studies, also including sampling of native mammals, are required on larger sample sizes and from wider geographic areas, to determine the significance of these findings, including the public health importance of Cryptosporidium spp. from rodents.     

Aspects of the life history of Cercopithifilaria johnstoni (Nematoda:Filarioidea)

and 1988. Aspects of the life history of Cercopithifilaria johnstoni (Nematoda:Filarioidea). International Journal for Parasitology 18: 1087–1092. Cercopithifilaria johnstoni (Nematoda:Filarioidea) occurs in the subcutaneous connective tissues of a spectrum of native murid and marsupial hosts in Eastern Australia. Life cycle studies revealed that: (i) microfilaria occur in lymphatic capillaries and extravascular connective tissue of the dermis (= ‘skin-inhabiting’), (ii) ixodid ticks, particularly Ixodes trichosuri, are intermediate hosts in nature, (iii) development from microfilariae to infective third-stage larva occurs only while the tick is off the host, that is, during ecdysis from larva to nymph or from nymph to adult. Transmission of C. johnstoni in a wild population of bush rats (Rattus fuscipes) occurred in summer and winter, and was associated with peaks in the number of larval and/or nymphal stages of ticks on rats. C. johnstoni was transmitted experimentally to bandicoots (Isoodon macrourus, Perameles nasuta), bush rats and laboratory rats (R. norvegicus), indirectly by subcutaneous inoculation of third-stage larvae and directly by tick feeding. The prepatent period was approximately 3 months and the longest duration of microfilariae in the ‘ skin’ was more than 25 months. Dermal and ocular lesions were observed in R. fuscipes. The host-parasite relationship has the potential for development as an inexpensive and practical model for human onchocerciasis.     

Epidemic increase of Endophyllum osteospermi (Uredinales, Pucciniaceae) on Chrysanthemoides monilifera

The epidemiology of the rust fungus Endophyllum osteospermi was investigated. This rust fungus is considered to be a candidate biological control agent for Chrysanthemoides monilifera ssp. monilifera, which is an invasive alien weed of native vegetation in south-eastern Australia. Between 10 and 20 plants of C. monilifera were marked at each of five sites in the Western Cape Province of South Africa, where both organisms are native. The infection levels and number of witches' brooms were determined every 2 months over a 2-year period. Additionally, at three of these sites, the infection levels and number of witches' brooms of all bushes in the host population was determined annually over 4 years. The increase in number of witches' brooms per bush ranged between 0 and 282 within 1 year, with an average increase per bush of 28 (SE±4.8) in 1993, and 39 (SE±9.2) in 1994. The average r_s for all bushes during 1993 was 0.015 month^-1 (SE±0.0041, n=72) and 0.0098 month^-1 (SE±0.0073, n=43) during 1994. When host bushes that either died back or died during the course of each year were excluded, then the average r_s during 1993 was 0.023 month^-1 (SE±0.0048, n=45) and 0.0348 month^-1 (SE±0.0106, n=20) during 1994. Under suitable conditions in South Africa, E. osteospermi undergoes epidemic increase within its host plant's populations. This rust fungus should therefore be considered as a suitable candidate biological control agent for use in Australia against C. monilifera ssp. monilifera.     

Multi-species predator eradication within a predator-proof fence at Ka‘ena Point, Hawai‘i

Ka‘ena Point Natural Area Reserve on O‘ahu hosts one of the largest seabird colonies in the main Hawaiian Islands and supports three species of endangered plants. In order to stop chronic predation by invasive alien mammals on native species, a peninsula-style predator-proof fence was constructed around a 20-ha portion of the reserve in 2011. Multi-species predator removal efforts began upon fence completion; diphacinone poison in bait boxes spaced 25 m apart was used to remove black rats, house mice, and small Indian mongooses. House mice also were removed with multiple-catch live traps spaced 12.5 m apart. Feral cats were removed with padded leg-hold traps. Feral cats and mongooses were eradicated in 1 month, black rats were eradicated in 2.5 months, and house mice were eradicated in about 9 months. Since eradication, incursions of cats and mongoose have been rare (1/7.2 months), but incursion frequency has been higher for black rats (1/56 days) and house mice (1/36–47 days). Buffer predator control was conducted to limit predator access and prevent reinvasion around the fence ends along the shoreline. Even with the high initial fence cost and ongoing predator incursion management, this method is expected to become more cost effective than previous predator control efforts after 16 years. Record numbers of Wedge-tailed shearwaters and Laysan albatrosses have fledged from the reserve after predator eradication, and regeneration of native plants and invertebrates is being observed. With careful planning and persistence, predator fences can be a cost-effective method of protecting natural resources, and multiple species of predators can be eradicated with traps and first-generation anti-coagulents.     

Are introduced black rats (Rattus rattus) a functional replacement for mycophagous native rodents in fragmented forests?

The fungal diet of the introduced black rat (Rattus rattus) was examined in a fragmented forest in northeastern New South Wales, Australia, to determine whether this species was consuming and dispersing the spores of native truffles. Because of the absence of native rodents at the site, the diet of the swamp wallaby (Wallabia bicolor), a known mycophagist in the region, was examined simultaneously as a benchmark against which to compare fungal consumption by black rats. All 19 scats collected from black rats contained fungal spores, while 29 of the 34 swamp wallaby scats contained fungal spores. Most spores were from hypogeous (‘truffle-like’) fungal species, although both black rats and swamp wallabies each consumed a few epigeous (‘mushroom-like’) taxa. While rat and wallaby diets contained many of the same taxa, their diets were significantly different in terms of the fungal taxa that comprised each sample. Our results suggest that black rats might perform an important spore dispersal role in degraded and fragmented landscapes where native rodents have been extirpated, and might complement the dispersal role played by larger mycophagous mammals like swamp wallabies.     

Potential spread of introduced black rat (Rattus rattus) parasites to endemic deer mice (Peromyscus maniculatus) on the California Channel Islands

Introduced species have the potential to outperform natives in two primary ways: via increased rates of predation and competition, and via the introduction of new parasites against which native species often lack effective immune defences. To assess the extent to which invasive species' parasites spread to native hosts, we compared the composition of helminth parasites found in introduced black rat ( Rattus rattus ) and endemic deer mouse ( Peromyscus maniculatus ) populations on a subset of the California Channel Islands. Results suggest that the whipworm, Trichuris muris , may have spread from introduced black rats to endemic island deer mice and has continued to thrive in one island population where rats were recently eradicated. These results yield two important conservation messages: (1) although the parasites introduced with invasive species may be few, they should not be ignored as they can spread to native species, and (2) introduced parasites have the potential to remain in a system even after their founding host is extirpated. These findings underscore the importance of parasitological surveys in invasive species research and baseline data for ecosystems where exotic species are likely to invade.     

Shifts from native to invasive small mammals across gradients from tropical forest to urban habitat in Borneo

Urbanization has paved the way for the spread of commensal rodents at global scale. However, it is largely unknown how these species use tropical anthropogenic landscapes originally covered with forests and inhabited by diverse small mammal assemblages. We surveyed non-flying small mammals in various urban and suburban habitat types and adjacent forest in the tropical town of Kota Kinabalu in Borneo. We used occupancy and polynomial regression models to determine variation in species occurrences along gradients of land-use intensity. Müller’s sundamys (Sundamys muelleri) was the only native small mammal species found in urban and suburban landscapes with a continuous decrease in occurrence probability from forests to urban habitats. The invasive Asian black rat (Rattus rattus species complex) and the invasive Asian house shrew (Suncus murinus) had the highest occurrence probabilities in habitats of intermediate land-use intensity, but Asian black rats are also likely to occasionally invade forested habitats and occupied urban habitats in sympatry with the Norway rat (Rattus norvegicus). In urban and suburban habitats, fallow land possibly favoured the occurrence of S. muelleri and S. murinus. Other native small mammal species (Muridae, Sciuridae, Tupaiidae) were found only in forested areas. Our study shows that native small mammals found in forest are largely replaced by invasive species in urban and suburban habitats. Due to their occurrence in habitats of various land use intensities, S. muelleri and R. rattus comprise central links between forest wildlife and urban species, an association that is important to consider in studies of parasite and disease transmission dynamics.     

Analysis of lacI mutations in Big Blue transgenic mice subjected to parasite-induced inflammation.

Parasite infections have long been associated with specific types of human cancers. Schistosoma hematobium is an inducer of urinary bladder cancer, Helicobacter pylori is a gastric carcinogen, and hepatitis B virus and Opisthorchis viverrini are causative agents of hepatocellular carcinoma. Another liver fluke, Fasciola hepatica, has also been identified as a neoplastic risk agent, primarily in animals. We used F. hepatica-induced inflammation in mice to determine if the presence of an aggressive liver fluke could induce mutagenic events in mammalian tissue. This provides a perspective on the relationship between chronic inflammation and cancer and may be a model for future studies on this complex association. In previous studies using the Big Blue^® transgenic mouse assay, we demonstrated an increase in lacI mutations in liver cells harvested from mice harboring F. hepatica flukes when compared to uninfected control animals. In these studies, we report on the types of mutations associated with this parasite infection. The observed mutational spectrum roughly corresponded to the spectrum of spontaneous mutations in liver cells when compared to control (uninfected) animals. However, the spectrum of mutations from parasitized animals showed a significant increase in complex changes and multiple mutations (18.2%) when compared to what would be expected from control animals (2.8%).     

Helminths from introduced small mammals on Kerguelen, Crozet, and Amsterdam Islands (southern Indian Ocean).

Four monoxenous nematodes and 1 heteroxenous cestode were found in 4 species of introduced small mammals on isolated sub-Antarctic islands of the Indian Ocean. In the Kerguelen Archipelago, Syphacia obvelata, Passalurus ambiguus (Nematoda: Oxyuridae), and Rodentolepis straminea (Cestoda: Cyclophyllidae) were respectively found in the house mouse Mus musculus, the rabbit Oryctolagus cuniculus, and the black rat Rattus rattus. One accidental nematode, Trichostrongylus sp. (Nematoda: Trichostrongylidae), was also found in a black rat on Kerguelen. On Possession Island (Crozet Archipelago), R. straminea was present in the black rat. On Amsterdam Island, the brown rat R. norvegicus harbored 2 species, R. straminea and Nippostrongylus brasiliensis (Nematoda: Heligmonellidae). The small number of founder hosts and the depauperate terrestrial communities on these remote islands explain the low diversity in the helminth communities of these introduced mammals compared with continental populations.     

Combining population genomics and ecological niche modeling to assess taxon limits between Carex jemtlandica and C. lepidocarpa

Carex section Ceratocystis (Cyperaceae) is a group of recently evolved plant species, in which hybridization is frequent, introgression is documented, taxonomy is complex, and morphological boundaries are vague. Within this section, a unified taxonomic treatment of the Carex jemtlandica–Carex lepidocarpa species complex does not exist, and Norway may currently be the sole country accepting species rank for both. Carex jemtlandica is mainly confined to Fennoscandia and is thus a Fennoscandian conservation responsibility. This motivated us to test the principal hypothesis that both C. jemtlandica and C. lepidocarpa represent evolutionary significant units, and that both deserve their current recognition at species level. We investigated their evolutionary distinctiveness in Norway,using restriction site-associated DNA sequencing and ecological niche modeling. Our genomic results reveal two genetic clusters, largely corresponding to C. jemtlandica and C. lepidocarpa that also remain distinct in sympatry, despite clear indications of ongoing hybridization and introgression. The ecological niche modeling suggests that they occupy different environmental niches. Jointly, our results clearly show that C. jemtlandica and C. lepidocarpa represent separately evolving entities that should qualify recognition as evolutionary significant units. Given the high level of introgression compared to other hybridizing species pairs in Carex we recommend treating C. jemtlandica as a subspecies of C. lepidocarpa.     

Murine viruses in an island population of introduced house mice and endemic short-tailed mice in Western Australia

House mice (Mus domesticus) were recently introduced to Thevenard Island, off the northwest coast of Western Australia. This island is also habitat for an endangered native rodent, the short-tailed mouse (Leggadina lakedownensis). Concerns have been raised that house mice may pose a threat to L. lakedownensis both through competition and as a source of infection. To assess the threat to L. lakedownensis posed by viral pathogens from M. domesticus, a serological survey was conducted from 1994 to 1996 of both species for evidence of infection with 14 common murine viruses (mouse hepatitis virus, murine cytomegalovirus, lymphocytic choriomeningitis virus, ectromelia virus, mouse adenovirus strains FL and K87, minute virus of mice, mouse parvovirus, reovirus type 3, Sendai virus, Theiler's mouse encephalomyelitis virus, polyoma virus, pneumonia virus of mice, and encephalomyocarditis virus) and Mycoplasma pulmonis. Despite previous evidence that populations of free-living M. domesticus from various locations on the Australian mainland were infected with up to eight viruses, M. domesticus on Thevenard Island were seropositive only to murine cytomegalovirus (MCMV). Antibodies to MCMV were detected in this species at all times of sampling, although seroprevalence varied. Infectious MCMV could be isolated in culture of salivary gland homogenates from seropositive mice. In contrast, L. lakedownensis on Thevenard Island showed no serological evidence of infection with MCMV, any of the other murine viruses, or M. Pulmonis, and no virus could be isolated in culture from salivary gland homogenates. Although MCMV replicated to high titers in experimentally infected inbred BALB/c laboratory mice as expected, it did not replicate in the target organs of experimentally inoculated L. lakedownensis, indicating that the strict host specificity of MCMV may prevent its infection of L. lakedownensis. These results suggest that native mice on Thevenard Island are not at risk of MCMV infection from introduced house mice, and raise interesting questions about the possible selective survival of MCMV in small isolated populations of M. domesticus.     

The reproductive tract of the male spiny mouse (Acomys cahirinus) and coagulation studies with other species

The testes of the spiny mice showed asymmetry, the left being significantly heavier than the right (P = 0.025). Histological studies indicated that spermatozoa were first present in the testes of animals 35--45 days of age but the maturation of the accessory glands, especially the lateral prostates and coagulating glands, occurred later. The highest fructose concentration in the adult was in the lateral prostates (126.97 +/- 22.23 mg fructose/100 g, n = 5) and coagulating glands (99.38 +/- 17.65 mg fructose/100 g gland weight, n = 5). Coagulation tests of mixtures of extracts of seminal vesicles and coagulating glands from spiny mice and rats indicated that the vesiculase of the spiny mouse was active on rat substrates and vice versa. Cross-reactions of extracts of house mouse (Mus musculus), hamster (Mesocricetus auratus), gerbil (Meriones unguiculatus), and guinea-pig (Cavia porcellus) seminal vesicles (substrate) and coagulating glands (vesiculase) with those of rats and spiny mice showed that although the substrates of rat and spiny mouse were readily coagulated by vesiculase from all the other species, rat and spiny mouse vesiculase were not equally active on substrates of the other species.     

Enhanced liver cell mutations in trematode-infected Big Blue transgenic mice

Parasite infections in humans have long been associated with specific types of cancers. Schistosoma hematobium is a known inducer of urinary bladder cancer, Helicobacter pylori is a gastric carcinogen, and hepatitis B virus and Opisthorchis viverrini are causative agents of liver cell cancers. Another liver fluke, Fasciola hepatica, has also been identified as a neoplastic risk agent, primarily in animals. We used F. hepatica as a model agent to determine if the presence of an aggressive liver fluke could induce mutagenic events in mammalian tissue. Using the Big Blue^® transgenic mouse assay, we found a two-fold increase in lacI mutations in cells harvested from mice harboring F. hepatica worms when compared to uninfected control animals. These data indicate that biological infections can cause increased genetic damage in surrounding host tissue.     

Dietary niche differentiation among three species of invasive rodents (Rattus rattus, R. exulans, Mus musculus)

The diets of sympatric rodents partially define their realized niches. Identifying items in stomachs of introduced rodents helps determine rodents’ trophic positions and species most at risk of consumption. In the Hawaiian Islands, which lacked rodents prior to human arrival, three rodents (Rattus rattus or black rat, R. exulans or Pacific rat, Mus musculus or house mouse) commonly coexist in native habitats where they consume a wide range of plants and animals. These three rodent species were trapped in montane forest for 2.5 years; their stomach contents were analyzed to determine short-term diets (n = 12–95 indiv. per species), and isotopic fractions of δ¹⁵N and δ¹³C in their bone collagen were analyzed to further estimate their trophic positions (n = 11–20 indiv. per species). For all three species, >75 % of individuals had plants and >90 % had arthropods in their stomachs, and significant differences in mean relative abundances were found for food items in stomachs among all three rodents. Rodents may be dispersing some native and non-native seeds, including the highly invasive Clidemia hirta. Most identifiable arthropods in rodent stomachs were non-native, and no stomachs contained birds, snails, or lizards. The δ¹⁵N and δ¹³C signatures were consistent with trophic feeding differences revealed from stomach contents. Dietary niche differentiation by coexisting rodent species is evident in this forest, with Pacific rats being intermediate between the mostly carnivorous house mouse and the mostly herbivorous black rat; such findings can help forecast rodent impacts and direct management efforts in ecosystems where these invasive animals coexist.