Phylogeny of the genus Apodemus with a special emphasis on the subgenus Sylvaemus using the nuclear IRBP gene and two mitochondrial markers: cytochrome b and 12S rRNA

Phylogenetic relationships among 17 extant species of Murinae, with special reference to the genus Apodemus, were investigated using sequence data from the nuclear protein-coding gene IRBP (15 species) and the two mitochondrial genes cytochrome b and 12S rRNA (17 species). The analysis of the three genes does not resolve the relationships between Mus, Apodemus, and Rattus but separates Micromys from these three genera. The analysis of the two mitochondrial regions supported an association between Apodemus and Tokudaia and indicated that these two genera are more closely related to Mus than to Rattus or Micromys. Within Apodemus, the mitochondrial data sets indicated that 8 of the 9 species analyzed can be sorted into two main groups: an Apodemus group, with A. agrarius, semotus, and peninsulae, and a Sylvaemus group, with uralensis, flavicollis, alpicola, sylvaticus, and hermonensis. The position of Apodemus mystacinus is ambiguous and might be either included in Sylvaemus or considered a distinct subgenus, Karstomys, more closely related to Sylvaemus than to Apodemus. Estimation of the divergence time for these taxa suggests a separation between 7 and 8 My ago for the three groups (mystacinus and the two subgenera Apodemus and Sylvaemus). Within each subgenus, divergence times are between 5.4 and 6 My for Apodemus and between 2.2 and 3.5 My for Sylvaemus and mystacinus.     

Hyperconservation of the N-formyl peptide binding site of M3: evidence that M3 is an old eutherian molecule with conserved recognition of a pathogen-associated molecular pattern

The mouse MHC class I-b molecule H2-M3 has unique specificity for N-formyl peptides, derived from bacteria (and mitochondria), and is thus a pathogen-associated molecular pattern recognition receptor (PRR). To test whether M3 was selected for this PRR function, we studied M3 sequences from diverse murid species of murine genera Mus, Rattus, Apodemus, Diplothrix, Hybomys, Mastomys, and Tokudaia and of sigmodontine genera Sigmodon and PEROMYSCUS: We found that M3 is highly conserved, and the 10 residues coordinating the N-formyl group are almost invariant. The ratio of nonsynonymous and synonymous substitution rates suggests the Ag recognition site of M3, unlike the Ag recognition site of class I-a molecules, is under strong negative (purifying) selection and has been for at least 50-65 million years. Consistent with this, M3 alpha1alpha2 domains from Rattus norvegicus and Sigmodon hispidus and from the "null" allele H2-M3(b) specifically bound N-formyl peptides. The pattern of nucleotide substitution in M3 suggests M3 arose rapidly from murid I-a precursors by an evolutionary leap ("saltation"), perhaps involving intense selective pressure from bacterial pathogens. Alternatively, M3 arose more slowly but prior to the radiation of eutherian (placental) mammals. Older dates for the emergence of M3, and the accepted antiquity of CD1, suggest that primordial class I MHC molecules could have evolved originally as monomorphic PRR, presenting pathogen-associated molecular patterns. Such MHC PRR molecules could have been preadaptations for the evolution of acquired immunity during the early vertebrate radiation.     

Functional respiratory chain analyses in murid xenomitochondrial cybrids expose coevolutionary constraints of cytochrome b and nuclear subunits of complex III

The large number of extant Muridae species provides the opportunity of investigating functional limits of nuclear/mitochondrial respiratory chain (RC) subunit interactions by introducing mitochondrial genomes from progressively more divergent species into Mus musculus domesticus mtDNA-less (rho0) cells. We created a panel of such xenomitochondrial cybrids, using as mitochondrial donors cells from six murid species with divergence from M. m. domesticus estimated at 2 to 12 Myr before present. Species used were Mus spretus, Mus caroli, Mus dunni, Mus pahari, Otomys irroratus, and Rattus norvegicus. Parsimony analysis of partial mtDNA sequences showed agreement with previous molecular phylogenies, with the exception that Otomys did not nest within the murinae as suggested by some recent nuclear gene analyses. Cellular production of lactate, a sensitive indicator of decreased respiratory chain ATP production, correlated with divergence. Functional characterization of the chimeric RC complexes in isolated mitochondria using enzymological analyses demonstrated varying decreases in activities of complexes I, III, and IV, which have subunits encoded in both mitochondrial and nuclear genomes. Complex III showed a striking decline in electron transfer function in the most divergent xenocybrids, being greatly reduced in the Rattus xenocybrid and virtually absent in the Otomys xenocybrid. This suggests that nuclear subunits interacting with cytochrome b face the greatest constraints in the coevolution of murid RC subunits. We sequenced the cytochrome b gene from the species used to identify potential amino acid substitutions involved in such interactions. The greater sensitivity of complex III to xenocybrid dysfunction may result from the encoding of redox center apoproteins in both nuclear and mitochondrial genomes, a unique feature of this RC complex.     

Identification of novel rodent herpesviruses, including the first gammaherpesvirus of Mus musculus

Rodent herpesviruses such as murine cytomegalovirus (host, Mus musculus), rat cytomegalovirus (host, Rattus norvegicus), and murine gammaherpesvirus 68 (hosts, Apodemus species) are important tools for the experimental study of human herpesvirus diseases. However, alphaherpesviruses, roseoloviruses, and lymphocryptoviruses, as well as rhadinoviruses, that naturally infect Mus musculus (house mouse) and other Old World mice are unknown. To identify hitherto-unknown rodent-associated herpesviruses, we captured M. musculus, R. norvegicus, and 14 other rodent species in several locations in Germany, the United Kingdom, and Thailand. Samples of trigeminal ganglia, dorsal root ganglia, brains, spleens, and other organs, as well as blood, were analyzed with a degenerate panherpesvirus PCR targeting the DNA polymerase (DPOL) gene. Herpesvirus-positive samples were subjected to a second degenerate PCR targeting the glycoprotein B (gB) gene. The sequences located between the partial DPOL and gB sequences were amplified by long-distance PCR and sequenced, resulting in a contiguous sequence of approximately 3.5 kbp. By DPOL PCR, we detected 17 novel betaherpesviruses and 21 novel gammaherpesviruses but no alphaherpesvirus. Of these 38 novel herpesviruses, 14 were successfully analyzed by the complete bigenic approach. Most importantly, the first gammaherpesvirus of Mus musculus was discovered (Mus musculus rhadinovirus 1 [MmusRHV1]). This virus is a member of a novel group of rodent gammaherpesviruses, which is clearly distinct from murine herpesvirus 68-like rodent gammaherpesviruses. Multigenic phylogenetic analysis, using an 8-kbp locus, revealed that MmusRHV1 diverged from the other gammaherpesviruses soon after the evolutionary separation of Epstein-Barr virus-like lymphocryptoviruses from human herpesvirus 8-like rhadinoviruses and alcelaphine herpesvirus 1-like macaviruses.     

西藏农区鼠害调查初报

2007年4月和6月在西藏山南、日喀则地区及拉萨市农区进行鼠密度和鼠种调查.采用铗夜法,共捕鼠169只,隶属2科4属5种,分别为小家鼠、黄胸鼠、白尾松田鼠、藏仓鼠和长尾仓鼠.白尾松田鼠为该区农田优势种,其组成比例为93.44%;黄胸鼠为山南地区农舍区的优势种,组成比例为82.35%.在日喀则农舍区则只捕获有小家鼠.山南和日喀则地区4月农田的平均捕获率分别为2.06%和7.43%;农舍区的平均捕获率分别为9.77%和9.15%.6月山南地区农田的平均捕获率为5.37%,显著高于该地4月的捕获率.西藏农区的白尾松田鼠在4月开始进入繁殖期.     

Dating of divergences within the Rattus genus phylogeny using whole mitochondrial genomes

The timing and order of divergences within the genus Rattus have, to date, been quite speculative. In order to address these important issues we sequenced six new whole mitochondrial genomes from wild-caught specimens from four species, Rattus exulans, Rattus praetor, Rattus rattus and Rattus tanezumi. The only rat whole mitochondrial genomes available previously were all from Rattus norvegicus specimens. Our phylogenetic and dating analyses place the deepest divergence within Rattus at approximately 3.5 million years ago (Mya). This divergence separates the New Guinean endemic R. praetor lineage from the Asian lineages. Within the Asian/Island Southeast Asian clade R. norvegicus diverged earliest at approximately 2.9Mya. R. exulans and the ancestor of the sister species R. rattus and R. tanezumi subsequently diverged at approximately 2.2Mya, with R. rattus and R. tanezumi separating as recently as approximately 0.4Mya. Our results give both a better resolved species divergence order and diversification dates within Rattus than previous studies.     

Evolutionary systematics of the Indian mouse Mus famulus Bonhote, 1898: molecular (DNA/DNA hybridization and 12S rRNA sequences) and morphological evidence

The genus Mus encompasses 38 species of mice divided into four subgenera: Mus , Pyromys , Nannomys and Coelomys . Each of these four taxa is characterized by discrete morphological as well as biochemical traits. We used two different molecular approaches to determine the relationships between these subgenera: DNA/DNA hybridization and 12S rRNA mitochondrial sequences. We compared the resulting phylogenies from each method and with phylogenies derived from morphological data. The degree of resolution of each molecular approach is discussed. The two molecular studies indicate that Mus , Pyromys , Nannomys and Coelomys are clearly distinct monophyletic groups, as previously indicated by morphological data and other biochemical and molecular approaches. There is one divergence between previous morphological and the molecular and morphological studies presented here: the position of the Indian species Mus famulus . This taxon, which was formerly included in the subgenus Coelomys , is demonstrated here to belong to the subgenus Mus. We also propose the following relationships within Mus sensu lato : Mus and Pyromys are the closest relatives, followed by Nannomys and Coelomys , whose relationships are still unclear. This arrangement is more robustly supported by DNA/DNA hybridization than by 12S rRNA data. A molecular time scale for the evolution within Mus sensu lato is proposed, using as a reference the Mus/Rattus divergence estimated by the fossil record at around 12 mya.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 137 , 385–401.     

Albumin evolution and phylogenetic relationships among Greek rodents of the families Arvicolidae and Muridae

The phylogenetic relationships of seven rodent species Microtus atticus, M. thomasi, M. epiroticus (family Arvicolidae) and Mus domesticus, Rattus norvegicus, Apodemus flavicollis and A. mystacinus (family Muridae) have been studied. In order to define these relationships we study the albumin evolution using the micro-complement fixation test (MC'F). No phylogenetic (immunological) distance between M. atticus and M. thomasi was found, a fact which confirms from the biochemical point of view the opinion that the former taxon is a synonym of the latter one. A molecular time scale relating MC'F immunological distances and geological time was established based on the assumption of a rate of 100 amino acid substitutions per–20 million years. The time of divergence between M. epiroticus and M. thomasi was estimated to be 0.5–0.6 million years ago (Pleistocene). Such a recent divergence corroborates the opinion based on morphological and protein electrophoretic criteria according to which Terricola (formerly Pitymys ) must be considered as a subgenus of the genus Microtus and not as a distinct genus Pitymys , as previously had been accepted. Apodemus flavicollis and A. mystacinus were separated about 0.65–0.8 million years ago (Pleistocene). The Rattus norvegicus lineage was separated–12.5 million years ago (end of Miocene), shortly before the Mus and Apodemus divergence. Our data indicate that the common ancestor of Arvicolidae and Muridae lived–25 million years ago (early Miocene). All these results are in agreement with paleontological and some recent DNA-DNA hybridization and electrophoretic data.     

Multiple hybridization events between Drosophila simulans and Drosophila mauritiana are supported by mtDNA introgression

The study of speciation has advanced considerably in the last decades because of the increased application of molecular tools. In particular, the quantification of gene flow between recently diverged species could be addressed. Drosophila simulans and Drosophila mauritiana diverged, probably allopatrically, from a common ancestor approximately 250,000 years ago. However, these species share one mitochondrial DNA (mtDNA) haplotype indicative of a recent episode of introgression. To study the extent of gene flow between these species, we took advantage of a large sample of D. mauritiana and employed a range of different markers, i.e. nuclear and mitochondrial sequences, and microsatellites. This allowed us to detect two new mtDNA haplotypes (MAU3 and MAU4). These haplotypes diverged quite recently from haplotypes of the siII group present in cosmopolitan populations of D. simulans. The mean divergence time of the most diverged haplotype (MAU4) is approximately 127,000 years, which is more than 100,000 years before the assumed speciation time. Interestingly, we also found some evidence for gene flow at the nuclear level because an excess of putatively neutral loci shows significantly reduced differentiation between D. simulans and D. mauritiana. Our results suggest that these species are exchanging genes more frequently than previously thought.     

乌鲁木齐市小型兽类群落特征与分布格局初步研究

根据2002年7月的调查资料对乌鲁木齐市养禽场鼠类群落进行了研究。结果表明：该地区分布的鼠类可划分为2种群落,(1)褐家鼠 小家鼠群落;(2)小家鼠 灰仓鼠群落。同时对鼠类群落的多样性,均匀度等进行了初步研究。     

A pronounced evolutionary shift of the pseudoautosomal region boundary in house mice

The pseudoautosomal region (PAR) is essential for the accurate pairing and segregation of the X and Y chromosomes during meiosis. Despite its functional significance, the PAR shows substantial evolutionary divergence in structure and sequence between mammalian species. An instructive example of PAR evolution is the house mouse Mus musculus domesticus (represented by the C57BL/6J strain), which has the smallest PAR among those that have been mapped. In C57BL/6J, the PAR boundary is located just ~700 kb from the distal end of the X chromosome, whereas the boundary is found at a more proximal position in Mus spretus, a species that diverged from house mice 2-4 million years ago. In this study we used a combination of genetic and physical mapping to document a pronounced shift in the PAR boundary in a second house mouse subspecies, Mus musculus castaneus (represented by the CAST/EiJ strain), ~430 kb proximal of the M. m. domesticus boundary. We demonstrate molecular evolutionary consequences of this shift, including a marked lineage-specific increase in sequence divergence within Mid1, a gene that resides entirely within the M. m. castaneus PAR but straddles the boundary in other subspecies. Our results extend observations of structural divergence in the PAR to closely related subspecies, pointing to major evolutionary changes in this functionally important genomic region over a short time period.     

Ribosomal DNA variation within and between species of rodents, with emphasis on the genus Onychomys

Patterns of restriction-endonuclease site and length variation at the nuclear rDNA locus (18S + 28S rRNA gene complex) were examined in rodents. Of the 164 restriction sites mapped for seven species, 22 were conserved (mapping to the 18S, 28S, and 5.8S genes and ITS1) in all three Onychomys species as well as in Mus musculus and in three closely related peromyscine rodents, Peromyscus boylii, P. eremicus, and Reithrodontomys megalotis. The nontranscribed spacer (NTS) region revealed most of the variation among these taxa, with the patterns of variation grouping into the following categories, (1) intraindividual variation revealing as many as four site-specific repeat types within an individual, (2) intraspecific and interspecific site variation confined to the NTS, and (3) length variation in both the transcribed and NTS regions. Length variation in the 28S rRNA gene was also examined in 17 additional rodent species, and most size differences mapped to the divergent domain, D8, found in sequence comparisons between Mus and Rattus. The systematic implications of rDNA variation are discussed using the perspective gained from these rodent comparisons.     

Confrontation of morphological and molecular data: the Praomys group (Rodentia, Murinae) as a case of adaptive convergences and morphological stasis

Phylogenetic relationships in a group of 21 African rodent species designated as the Praomys group (Murinae) were investigated using morphological characters and sequence data from the complete mitochondrial cytochrome b gene and nuclear IRBP gene fragment (840bp). The molecular results confirm the monophyly of the Praomys group, including the species Malacomys verschureni, while the other Malacomys species appear very divergent. The basal relationships within the Praomys group are poorly resolved, suggesting a rapid radiation at about 7-9 million years ago based on genetic divergence rates calibrated from the fossil record. Discrepancies between molecular and morphological results probably reflect of numerous convergences as well as variations in the rates of morphological evolution among lineages. Reconstructions of the ancestral character states suggest a savannah origin for the Praomys group, along with some morphological traits conserved by stasis in savannah taxa. At the same time, forest taxa seem to be characterized by an accelerated morphological evolution, with acquisition of convergent adaptive characters.     

Cross-transmission of Eimeria spp. (Protozoa, Apicomplexa) of rodents--a review

A total of 169 cross-transmission attempts has been made with 44 (11.8%) of the 372 named species of Eimeria of rodents. Of these, 161 were rodent-to-rodent, 6 rodent-to-lagomorph, and 1 each rodent-to-carnivore and rodent-to-bird. None of the last three categories was successful. In the rodent-to-rodent combinations, 39 (80%) of the 49 attempts to transmit a coccidian species from one rodent species to another of the same genus were successful, and only 14 (12.5%) of the 112 attempts to transmit a coccidium to a rodent of a different genus were successful. Eight of the successful attempts were with E. chinchillae, which was the only truly euryxenous species of Eimeria in the group. Two successful attempts were between between the closely related rodent genera Spermophilus and Cynomys, and two were both of E. separata from Rattus norvegicus to some genetic strains but not to others of Mus musculus. One attempt with E. vermiformis from Mus musculus to Rattus norvegicus required treatment of the rat with the immunosuppressant dexamethasone to succeed. More cross-transmission studies are needed to determine the host spectra of the species of Eimeria and other coccidian genera, and to determine the roles of genetics and immunosuppression in their transmission.     

Cross-Transmission of Eimeria spp. (Protozoa, Apicomplexa) of Rodents—A Review

A total of 169 cross-transmission attempts has been made with 44 (11.8%) of the 372 named species of Eimeria of rodents. Of these, 161 were rodent-to-rodent, 6 rodent-to-lagomorph, and 1 each rodent-to-carnivore and rodent-to-bird. None of the last three categories was successful. In the rodent-to-rodent combinations, 39 (80%) of the 49 attempts to transmit a coccidian species from one rodent species to another of the same genus were successful, and only 14 (12.5%) of the 112 attempts to transmit a coccidium to a rodent of a different genus were successful. Eight of the successful attempts were with E. chinchillae , which was the only truly euryxenous species of Eimeria in the group. Two successful attempts were between the closely related rodent genera Spermophilus and Cynomys , and two were both of E. separata from Rattus norvegicus to some genetic strains but not to others of Mus musculus. One attempt with E. vermiformis from Mus musculus to Rattus norvegicus required treatment of the rat with the immunosuppressant dexamethasone to succeed. More cross-transmission studies are needed to determine the host-spectra of the species of Eimeria and other coccidian genera, and to determine the roles of genetics and immunosuppression in their transmission.     

Adaptive functional divergence among triplicated alpha-globin genes in rodents

The functional divergence of duplicated genes is thought to play an important role in the evolution of new developmental and physiological pathways, but the role of positive selection in driving this process remains controversial. The objective of this study was to test whether amino acid differences among triplicated alpha-globin paralogs of the Norway rat (Rattus norvegicus) and the deer mouse (Peromyscus maniculatus) are attributable to a relaxation of purifying selection or to a history of positive selection that has adapted the gene products to new or modified physiological tasks. In each rodent species, the two paralogs at the 5'-end of the alpha-globin gene cluster (HBA-T1 and HBA-T2) are evolving in concert and are therefore identical or nearly identical in sequence. However, in each case, the HBA-T1 and HBA-T2 paralogs are distinguished from the third paralog at the 3'-end of the gene cluster (HBA-T3) by multiple amino acid substitutions. An analysis of genomic sequence data from several rodent species revealed that the HBA-T3 genes of Rattus and Peromyscus originated via independent, lineage-specific duplication events. In the independently derived HBA-T3 genes of both species, a likelihood analysis based on a codon-substitution model revealed that accelerated rates of amino acid substitution are attributable to positive directional selection, not to a relaxation of purifying selection. As a result of functional divergence among the triplicated alpha-globin genes in Rattus and Peromyscus, the red blood cells of both rodent species contain a mixture of functionally distinct alpha-chain hemoglobin isoforms that are predicted to have different oxygen-binding affinities. In P. maniculatus, a species that is able to sustain physiological function under conditions of chronic hypoxia at high altitude, the coexpression of distinct hemoglobin isoforms with graded oxygen affinities is expected to broaden the permissible range of arterial oxygen tensions for pulmonary/tissue oxygen transport.     

The involucrin genes of the mouse and the rat: study of their shared repeats

The involucrin genes of the mouse (Mus musculus) and the rat (Rattus norvegicus) have been cloned and sequenced. The coding region of each gene contains, at site P, a segment of repeats homologous to that of other nonanthropoid mammals. In contrast to the repeats of species belonging to different mammalian orders, many individual repeats of the mouse and the rat can be matched. Both before and after the divergence of the two species, these repeats have been the site of systematic alterations in nucleotide sequence. One of the alterations is the correction of nucleotides of one repeat by those of another. Corrected nucleotides may be closely linked to flanking nucleotides that are uncorrected; the systematic correction process therefore appears to be due to gene conversion. There is a stretch of 18 reiterated CAGs in the segment of repeats of the Mus gene; most of these reiterations were introduced recently, supporting the idea that the gene was generated originally from poly CAG. An antiserum to a synthetic peptide encoded by the segment of repeats of the Mus gene reveals differentiation- specific expression of the gene in the epidermis.      

洞庭平原农房家鼠的栖息格局

洞庭平原农房主要有褐家鼠、小家鼠、黄胸鼠。鼠密度最高为厨房, 其次为杂屋, 卧室和堂屋较低。房屋的不同地面对鼠密度与鼠种组成都有影响, 地面用水泥硬化后房间的鼠密度明显低于普通未硬化泥土地面,硬化地面以小家鼠为主, 泥土地面以褐家鼠占优势。不同结构房屋的鼠密度亦有差异, 砖混结构的鼠密度明显低于土木结构; 其鼠种组成也不同, 砖混结构的房屋小家鼠占优势, 土木结构房屋则褐家鼠与小家鼠比例相当。房屋地面的硬化对害鼠的影响比房屋结构的改变要大。综合来看, 鼠密度最高的房间为泥土地面+ 土木结构的厨房; 鼠密度最低为硬化地面+ 砖混结构的堂屋。通常, 在家具等物品摆放整洁、食物与水源少的硬化地面的房间鼠密度较低, 且以小家鼠占优势; 在杂物堆放较多、脏乱差的泥土地面房间的鼠密度较高。因此, 在防治中应根据不同的环境条件、不同的鼠密度和优势鼠种, 采取相应的防治对策, 以取得最佳防治效果。     

Genetic characterization of Okinawan black rats showing coat color polymorphisms of white spotting and melanism

We examined pelage color variation in wild populations of black rats (the Rattus rattus species complex) in the Yambaru forest area, northern Okinawa Island, Ryukyu Archipelago, Japan. Our field study revealed that 8.7% (38/438) and 0.2% (4/2500) of rats exhibited two types of coat color: white spotting and melanism, respectively. Using 34 representative animals, the phylogeography of the population was inferred using a nuclear gene marker, i.e., sequences (954 bp) of the melanocortin-1 receptor (Mc1r) gene responsible for the melanistic form in black rats. Four sequences from Okinawa were characterized as R. tanezumi, the Asian strain of black rat. Notably, neither of the phenotypic characters of white spotting or melanism was associated with the Mc1r haplotypes. Analysis of mitochondrial cytochrome b (Cytb) sequences (1140 bp) revealed that four haplotypes recovered from Okinawa clustered with the clade of R. tanezumi and differed by one or more bases from haplotypes at other localities in Japan and Asian countries. Thus, both variants may have arisen in the native rat population of Okinawa without interaction with the lineage of R. rattus, which exhibits a worldwide distribution and displays such coat color variants. The Yambaru population of black rats has thus experienced its own evolutionary history in allopatry for a substantial period of time (e.g., 10,000 years), which has preserved valuable genetic polymorphisms and will be useful for assessing the ecological consequences of genetic variation in natural populations.     

Evolution of the FAD2-1 fatty acid desaturase 5' UTR intron and the molecular systematics of Gossypium (Malvaceae)

The FAD2-1 microsomal omega-6 desaturase gene contains a large intron ( approximately 1133 bp [base pairs]) in the 5' untranslated region that may participate in gene regulation and, in GOSSYPIUM:, is evolving at an evolutionary rate useful for elucidating recently diverged lineages. FAD2-1 is single copy in diploid GOSSYPIUM: species, and two orthologs are present in the allotetraploid species. Among the diploid species, the D-genome FAD2-1 introns have accumulated substitutions 1.4-1.8 times faster than the A-genome introns. In the tetraploids, the difference between the D-subgenome introns and their A-subgenome orthologs is even greater. The substitution rate of the intron in the D-genome diploid G. gossypioides more closely approximates that of the A genome than other D genome species, highlighting its unique evolutionary history. However, phylogenetic analyses support G. raimondii as the closest living relative of the D-subgenome donor. The Australian K-genome species diverged 8-16 million years ago into two clades. One clade comprises the sporadically distributed, erect to suberect coastal species; a second clade comprises the more widely spread, prostrate, inland species. A comparison of published gene trees to the FAD2-1 intron topology suggests that G. bickii arose from an early divergence, but that it carries a G. australe-like rDNA captured via a previously undetected hybridization event.