Cytonuclear discordance among Southeast Asian black rats (Rattus rattus complex)

Black rats are major invasive vertebrate pests with severe ecological, economic and health impacts. Remarkably, their evolutionary history has received little attention, and there is no firm agreement on how many species should be recognized within the black rat complex. This species complex is native to India and Southeast Asia. According to current taxonomic classification, there are three taxa living in sympatry in several parts of Thailand, Cambodia and Lao People's Democratic Republic, where this study was conducted: two accepted species (Rattus tanezumi, Rattus sakeratensis) and an additional mitochondrial lineage of unclear taxonomic status referred to here as ‘Rattus R3’. We used extensive sampling, morphological data and diverse genetic markers differing in rates of evolution and parental inheritance (two mitochondrial DNA genes, one nuclear gene and eight microsatellite loci) to assess the reproductive isolation of these three taxa. Two close Asian relatives, Rattus argentiventer and Rattus exulans, were also included in the genetic analyses. Genetic analyses revealed discordance between the mitochondrial and nuclear data. Mitochondrial phylogeny studies identified three reciprocally monophyletic clades in the black rat complex. However, studies of the phylogeny of the nuclear exon interphotoreceptor retinoid‐binding protein gene and clustering and assignation analyses with eight microsatellites failed to separate R. tanezumi and R3. Morphometric analyses were consistent with nuclear data. The incongruence between mitochondrial and nuclear (and morphological) data rendered R. tanezumi/R3 paraphyletic for mitochondrial lineages with respect to R. sakeratensis. Various evolutionary processes, such as shared ancestral polymorphism and incomplete lineage sorting or hybridization with massive mitochondrial introgression between species, may account for this unusual genetic pattern in mammals.     

Sugar preferences of "black rats", Rattus rattus L.

The choice of "black rats", Rattus rattus L ., for common sugars - sucrose, jaggery, glucose, fructose and lactose are described. In laboratory colonies, the sugars were preferred in the order glucose > sucrose > jaggery > fructose > lactose; but in another sequence - sucrose > jaggery > glucose > fructose > lactose, in the free living colonies. The discrepancy is accounted for by the characteristics of sugars which influence consummatory behaviour in the two situations.     

Frequency of chromosome polymorphism in Rattus rattus collected in Japan

This paper deals with a population survey of chromosome polymorphism of Rattus rattus collected in Japan and the results of their test crosses. All the animals had diploid 42 chromosomes, but three chromosome pairs, Nos. 1, 9 and 13, were polymorphic in respect to acro- and subtelocentric chromosomes. Frequency of No. 1 chromosome polymorphism in 453 rats collected in 19 localities showed 343 rats (75.5%) with acrocentric homomorphic pair (A/A), 90 (19.9%) with aerocentric and subtelocentric heteromorphic pair (A/S) and the remaining 20 (4.4%) with subtelocentric homomorphic pair (S/S). All animals collected in northern and northwestern Japan showed only the A/A pair, while those collected in southern and southeastern Japan showed A/A, A/S and S/S polymorphism. The latter group was also classified into 3 populations (east, southeast and south) by the different frequency of the subtelocentric chromosome. Progeny tests revealed that segregation of A/A, A/S and S/S types from F₁ hybrids between various chromosome combinations was not significantly different from the theoretical one. However, the number of animals with A/S pair was always slightly higher than the other two types, while those with S/S pair slightly fewer. Local differences of the chromosome polymorphism in Japan were considered due to the result of migration and selection of the rats with S/S chromosome type.Contribution No. 817 from the National Institute of Genetics, Japan. Supported in part by a grant-in-aid from the Ministry of Education of Japan (Scientific Expedition in 1968, and No. 8801 in 1969).     

Preliminary genetic characterization of two lineages of black rats (Rattus rattus sensu lato) in Japan, with evidence for introgression at several localities

We conducted a pilot survey of genetic diversity among 37 karyotyped individuals of the black rat Rattus rattus (sensu lato) from six localities on the Japanese Islands, using complete gene sequences of mitochondrial cytochrome b (cyt b) and nuclear interphotoreceptor retinoid binding protein (IRBP). Our sampling included two previously documented karyotypic groups: 'Oceanian' with 2n = 38 and 'Asian' with 2n = 42. Cyt b sequences for most individuals clustered according to their karyotypic groups, with an average between-group divergence of 3.8%. One exception was that individuals from Kagoshima (Kyushu Island) showed 'Asian' karyotypes combined with a cyt b haplotype that differed by a single nucleotide substitution from the haplotype of the 'Oceanian' karyotypic group. Six IRBP haplotypes were identified. They belonged to three distinct IRBP lineages (I-III), with an average inter-lineage divergence of 1%. Among homozygous individuals, these lineages showed good association with the karyotypic groups: IRBP lineage I occurred only with 'Oceanian' karyotypes, while IRBP lineages II and III both occurred with 'Asian' karyotypes. Individuals from Kagoshima all possessed IRBP of 'Asian' lineages, despite the presence of an 'Oceanian' mitochondrial type. The Chichijima population (Ogasawara Islands) featured exclusively 'Asian' karyotypes and cyt b sequences, but various combinations of all three IRBP lineages. The Kagoshima and Chichijima populations thus provide strong evidence of viable hybridization and genetic introgression between the two karyotypic groups, but with variable genetic outcomes. Our results demonstrate the potential of combined analysis of karyotypes and mitochondrial and nuclear gene sequences to elucidate the complex dispersal and population history of the black rat.     

Karyotypes and serum transferrin patterns of hybrids between Asian and Oceanian black rats,Rattus rattus

Karyotypes and serum transferrin patterns were examined in Asian and Oceanian black rats (R. rattus). Japanese R. r. tanezumi and Malayan R. r. diardii had 2n=42, but Australian and New Guinea R. r. rattus showed 2n=38 chromosomes. F₁ hybrids between Japanese and Australian rats and Malayan and New Guinea rats had 2n=40 chromosomes which consists of the two genomes of both parents. Although various matings between the F₁ hybrids were made, only one F₂ male rat with 2n=39 chromosomes was obtained. The F₁ hybrids seem to be semisterile. Parental transferrin phenotypes were TfR in Japanese rats and TfCD in Oceanian rats. F₁ hybrids examined showed TfRD in both male and female and one F₂ hybrid had TfR type transferrin. Based on the above investigations, it is suggested that Asian and Oceanian black rats are geographically isolated and evolved different chromosomal and serum transferrin characteristics, but the sexual isolation of the two groups is incomplete at the present time.Contribution No. 826 from the National Institute of Genetics, Japan. Supported by a grant-in-aid from the Ministry of Education of Japan (Scientific Expedition in 1968, No. 8801 in 1969 and No. 9001 in 1970).     

Cytogenetical survey of black rats,Rattus rattus,in southwest and Central Asia,with special regard to the evolutional relationship between three geographical types

A chromosome survey of the black rat, Rattus rattus, was made from animals collected at different localities in Southwest and Central Asia. Asian type black rats (2 n=42) were distributed in northern India, northern Pakistan, while the Oceanian type rats (2 n=38) were found in southern India, southern Pakistan and Central Asia. A border line of distribution of rats with Asian and Oceanian types can be drawn dividing India and Pakistan into northern and southern parts. A hybrid type between Asian and Oceanian types was found in Karachi, Pakistan. Rats with 40 chromosomes, probably a transient type from Asian to Oceanian type, were found in Sri Lanka (Ceylon). It is suggested that these three geographic variants have developed via sequential events of Robertsonian fusion of acrocentric chromosomes in Asian type black rats. This fusion probably took place somewhere in southern India. The Oceanian type black rats that thus developed in southern India migrated widely to the rest of the world through Central Asia and Europe accompanying the movement of mankind.     

Population recovery of alien black rats Rattus rattus: A test of reinvasion theory

Reinvasion of pest animals after incomplete control is a major challenge for invasive species management, yet little is known about the behavioural and demographic categories of reinvaders or the mechanisms that drive population‐level responses to control. To understand the fine‐scale mechanisms of reinvasion, we examined changes in demography, movements and activity patterns of reinvading alien black rats Rattus rattus in the short (4 weeks) and longer term (3 months) following localised experimental pest removal. Using recovery and invasion theory, we tested three hypothesised mechanisms of reinvasion: the ‘in situ effect’, the ‘trickle effect’ and the ‘vacuum effect’. We created space for reinvasion by removing black rats from the core of replicate 1‐ha plots (short‐term experiment) and later by removing animals from the entire plot (longer‐term experiment). Reinvaders were characterised as dispersing juveniles, floaters or neighbours. Radio‐tracking quantified home range changes for adjacent resident animals (short‐term experiment only). In the short term, there was no net influx of rats after targeted removal. Radio‐tracked residents’ movements were highly variable and displayed no directional changes after nearby conspecifics were removed. However, in the longer term, removal led to slow population recovery through a mix of reinvading floaters, dispersing juveniles and shifting residents. These responses best support a hypothesis of reinvasion through a trickle effect, with rats being extremely mobile and having a high degree of population turnover, even in untreated sites. Our findings provide the first test of reinvasion theory at a small scale, demonstrating the importance of understanding the differing categories of reinvaders and mechanisms of reinvasion after population control. These mechanisms drive the rate of population recovery and, in turn, should help determine which strategy of pest control should be used, and the frequency with which they are implemented, in order to slow the recovery of pest 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.     

Chromosomal polymorphism in Rattus rattus (L.) collected in Kusudomari and Misima

Summary Chromosomes of Rattus rattus (L.), collected in Kusudomari (Nagasaki) and Misima (Sizuoka) were examined. The karyotype revealed a remarkable heteromorphism in chromosome no. 1. The homozygotic, i.e. standard type, was characterized by 13 pairs of telocentric and 7 pairs of metacentric chromosomes. Chromosome pair no. 1 was telocentric. X and Y chromosomes were also telocentrics. 18.4 per cent of rats from Kusudomari and 40 per cent from Misima showed heteromorphic pair in chromosome no. 1. One chromosome of the heteromorphic pair is conspicuous by the subtelocentric centromere. Total length of the telocentric chromosome of no. 1 is almost the same as of its subtelocentric partner. These facts indicate that the subtelocentric no. 1 chromosome might have arisen by a centromeric inversion of the telocentric chromosome. Individuals homozygous for the subtelocentric no. 1 chromosome could not be found in either population. The difference in the frequency of the dimorphics collected in Kusudomari and Misima was statistically significant. Possible causes of the difference are discussed.Dedicated to Professor H. Bauer on the occasion of his sixtieth birthday. — Contributions from the National Institute of Genetics, Misima, Japan, No. 533     

Frugivory by introduced black rats (Rattus rattus) promotes dispersal of invasive plant seeds

Oceanic islands have been colonized by numerous non-native and invasive plants and animals. An understanding of the degree to which introduced rats (Rattus spp.) may be spreading or destroying seeds of invasive plants can improve our knowledge of plant-animal interactions, and assist efforts to control invasive species. Feeding trials in which fruits and seeds were offered to wild-caught rats were used to assess the effects of the most common rat, the black rat (R. rattus), on 25 of the most problematic invasive plant species in the Hawaiian Islands. Rats ate pericarps (fruit tissues) and seeds of most species, and the impacts on these plants ranged from potential dispersal of small-seeded (≤1.5 mm length) species via gut passage (e.g., Clidemia hirta, Buddleia asiatica, Ficus microcarpa, Miconia calvescens, Rubus rosifolius) to predation where <15% of the seeds survived (e.g., Bischofia javanica, Casuarina equisetifolia, Prosopis pallida, Setaria palmifolia). Rats consumed proportionally more seed mass of the smaller fruits and seeds than the larger ones, but fruit and seed size did not predict seed survival following rat interactions. Although invasive rat control efforts focus on native species protection, non-native plant species, especially those with small seeds that may pass internally through rats, also deserve rat control in order to help limit the spread of such seeds. Black rats may be facilitating the spread of many of the most problematic invasive plants through frugivory and seed dispersal in Hawaii and in other ecosystems where rats and plants have been introduced.     

Tooth wear patterns in black rats (Rattus rattus) of Madagascar differ more in relation to human impact than to differences in natural habitats

Dietary characteristics and environmental variables are important selective factors directing ecological diversification in rodents. On Madagascar, the introductions and spread of the commensal black rat (Rattus rattus) can be seen as example cases to study dietary niche occupation and dietary adaptation in an insular environment. We investigate how tooth wear as a measure of dietary adaptation of black rats differs between four distinct habitats (village, manioc fields, spiny forest, and rainforest) with different dietary resources. We use the 3D surface texture analysis (3DST, using 30 parameters according to ISO 25178) as a measure of dietary abrasiveness. 3DST is applied on the occlusal surface of the upper first molar of 37 black rat specimens. The rainforest sample displays less rough and less voluminous surface textures compared to the village samples as indicated by smaller values for height parameters (Sa, Sp, Sq), inverse areal material ratio (Smc), and volume parameters (Vm, Vmc, Vmp, Vv, and Vvc). We therefore rank sampling areas from highest to lowest abrasiveness (village>manioc fields/spiny forest>rainforest). The rats from villages and rainforest differ to such an extent that one could have interpreted them to belong to different species. This indicates a high degree of variability in terms of ingesta abrasiveness. Furthermore, the pronounced difference between rats from human habitations compared to rats from associated fields or natural vegetation is interpreted to clearly indicate shifts in dietary niche occupation in relation to human impact.     

Frequencies of chromosome polymorphism in pairs no. 1, 9, and 13 in three geographical variants of black rats,Rattus rattus

Frequencies of the acrocentric and subtelocentric polymorphism in pairs no. 1, 9 and 13 chromosomes have been examined in 358 black rats, Rattus rattus, distributed over several countries of Asia, Australia and United States. The black rats are divided into three geographical types by the different chromosome numbers, such as Asian (2n=42), Ceylon (2n=40) and Oceanian types (2n=38). Pairs no. 13 polymorphism was found widely in these all types, but the pair no. 1 and 9 polymorphisms were found in only Asian type black rats. In the Asian type rats, however, those distributed in northern India and Pakistan showed always the subtelocentric pairs no. 1 and 9 like as those in Ceylon and Oceanian type black rats. This finding supports that the Ceylon and Oceanien type rats have developed in India or Pakistan from the Asian type. The present study also suggests that inversion of the pair no. 13 could have occurred in earlier period than those of the pairs no. 1 and 9.     

Molecular phylogeny of butterflies Parnassius glacialis and P. stubbendorfii at various localities in East Asia.

The phylogeny of butterflies, Parnassius stubbendorfii and P. glacialis, collected at various localities in the Japan archipelago and the eastern part of the Asian continent was analyzed using mitochondrial DNA sequences coding for NADH dehydrogenase subunit 5 (805 bp). The molecular phylogenetic trees revealed that P. glacialis and P. stubbendorfii diverged from a common ancestor, and then the populations inhabiting the Japan archipelago and the Asian continent diverged in each species. The reliability of these divergences was supported by high bootstrap values. The divergences within the Japan archipelago and within the Asian continent in each species were unclear because of low bootstrap values. The genetic distance and a rough time-estimation in the UPGMA tree suggest that the both populations of P. glacialis and P. stubbendorfii may have been isolated in the Japan archipelago at the early time (about 1.7-2.0 Mya) of the glacial period in the Pleistocene. The genetic distance between the Japanese and the continental subspecies may be large enough that they can be classified as different species, in comparison with the genetic distances among some other parnassian species.     

Invasion genetics of the introduced black rat (Rattus rattus) in Senegal,West Africa

An understanding of the evolutionary history and dynamics of invasive species is required for the construction of predictive models of future spread and the design of biological management measures. The black rat (Rattus rattus) is a major vertebrate invader with a worldwide distribution. Despite the severe ecological, economic and health impacts of this species, its evolutionary history has been little studied. We carried out extensive specimen sampling in Senegal, West Africa, and used microsatellite markers to describe the pattern and processes of invasion in this large continental area. The genetic data obtained were combined with historical knowledge concerning the presence of this species in Senegal. Data were analysed by a combination of Bayesian clustering and approximate Bayesian computation methods. The invasion pathways closely paralleled the history of human trade routes in Senegal. In several places, we detected the occurrence of multiple introductions from genetically different sources. Long‐distance migration between towns and villages was also observed. Our findings suggest that genetic bottlenecks and admixture have played a major role in shaping the genetics of invasive black rats. These two processes may generate genetic novelty and favour rapid evolution along the invasion pathways.     

Cotton rats (Sigmodon hispidus) and black rats (Rattus rattus) as possible reservoirs of Leishmania spp. in Lara State,Venezuela

A total of 519 wild animals belonging to eleven species were collected during a two year study in a cutaneous leishmaniasis endemic area in Venezuela (La Matica, Lara State). The animals were captured in home-made Tomahawk-like traps baited with maize, bananas or other available local fruits, and parasites were isolated from 27 specimens. Two different species were found naturally infected with flagellates, i.e., cotton rats (Sigmodon hispidus) and black rats (Rattus rattus). Characterization of the parasites using PCR, kDNA restriction pattern and hybridization with species-specific probes revealed the presence of Leishmania (L.) mexicana in three of the black rats and Leishmania (V.) braziliensis in two others. The latter species was also identified in the single positive specimen of S. hispidus. The results suggested both species of animals as possible reservoirs of Leishmania sp.     

Plasmodium berghei: malaria infection causes increased cardiac output in rats, Rattus rattus

Thirty-two 4-week-old male Wistar rats were infected with Plasmodium berghei malaria. On Days 12 through 14, blood volume, arterial blood pressure, right ventricular pressure, heart rate, cardiac output, stroke volume, hematocrit, and vascular resistances were determined. All of the cardiovascular parameters measured, with the exception of calculated pulmonary vascular resistance, changed progressively as the peripheral blood parasitemia increased. With a rising parasitemia, cardiac output increased, despite a reduced heart rate. The highest parasitemia of 63% was accompanied by a doubling of the normal cardiac output. The relationship between parasitemia and cardiac output can be described by the equation, cardiac output = (6.14) x % parasitemia + 452 ml/min/kg. The mean arterial blood pressure was lower than controls when parasitemia exceeded 20%, whereas systolic right ventricular pressure was elevated only at the highest parasitemias. When noninfected control rats were compared with those animals having parasitemias greater than 40%, in the infected animals, mean arterial pressure was 28% lower (P less than 0.01) and systolic right ventricular pressure rose by 21% (P less than 0.02). A 50% decline was observed in the total peripheral vascular resistance (P less than 0.01), although the pulmonary resistance was apparently unchanged. With P. berghei infection, there is also a marked anemia, an increase in plasma volume, and a 16% increase in blood volume (% body weight). It is concluded from these results that although the hemodynamic changes previously reported in the literature indicate that infection with malaria may result in focal blockages in microvessels and poor tissue perfusion, the total systemic effect, in the rat, is an increase in cardiac output secondary to a reduced peripheral resistance.(ABSTRACT TRUNCATED AT 250 WORDS)