Cyanogenesis variation in the Turnera sidoides L. polyploid complex (Turneraceae)

This study constitutes the first investigation of cyanogenesis variation within the Turnera sidoides autopolyploid complex. The qualitative and quantitative analysis carried out in 38 populations showed that the five subspecies of the complex are cyanogenic, although the levels of cyanide released varied both between and within subspecies. Between the subspecies, carnea , holosericea and integrifolia showed higher values of HCN released than subspecies pinnatifida and sidoides . Within subspecies, there is a tendency toward an increment of mean cyanide values with ploidy levels and different hypotheses are considered in order to explain this variation. The study of cyanogenesis in populations from different geographical areas cultivated under uniform greenhouse conditions showed that, in the widespread subspecies pinnatifida , HCN variation may also reflect the action of different selection pressures, since populations having dissimilar cyanide levels grow in different phytogeographical provinces with different climatic regimes. On the other hand, the fact that the variation of HCN released was greater between populations from different geographical areas than that observed between diploid and tetraploid populations of the same area provides additional evidence for the multiple origins of polyploids of the T. sidoides complex.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 85–94.     

Cytogeography of the South American Turnera sidoides L. complex (Turneraceae, Leiocarpae)

New chromosome number determinations are presented for 48 populations of five subspecies of Turnera sidoides. Chromosome counts in sspp. carnea, holosericea, integrifolia and sidoides confirm previous data while, in ssp. pinnatifida , the numbers 2n=2x = 14 and 2n = 6x = 42 are reported for the first time. The results show that polyploidy is a very frequent phenomenon. Populations are primarily tetraploids (2n = 4x = 28), while diploids (2n = 2x = 14), hexaploids (2n = 6x =42) and octoploids (2rc = 8x = 56) are less frequent. Diploid, tetraploid and hexaploid cytotypes occur in sspp. carnea and pinnatifida , whereas tetraploids and hexaploids are known in ssp. holosericea. Turnera sidoides ssp. integrifolia shows a polyploid series with ploidy levels from 2x to 8x. The ssp. sidoides appears to be uniformly tetraploid, with the exception of an isolated pentaploid (2rc = 5x = 35). On the basis of these counts and previous reports the geographical distribution of the cytotypes has been determined and related to climatic and ecological factors.     

How to look like a mallow: evidence of floral mimicry between Turneraceae and Malvaceae

Abundant, many-flowered plants represent reliable and rich food sources for animal pollinators, and may even sustain guilds of specialized pollinators. Contrastingly, rare plants need alternative strategies to ensure pollinators' visitation and faithfulness. Flower mimicry, i.e. the sharing of a similar flower colour and display pattern by different plant species, is a means by which a rare species can exploit a successful model and increase its pollination services. The relationship between two or more rewarding flower mimic species, or Müllerian mimicry, has been proposed as mutualistic, in contrast to the unilaterally beneficial Batesian floral mimicry. In this work, we show that two different geographical colour phenotypes of Turnera sidoides ssp. pinnatifida resemble co-flowering Malvaceae in colour as seen by bees' eyes, and that these pollinators do not distinguish between them when approaching flowers in choice tests. Main pollinators of T. sidoides are bees specialized for collecting pollen in Malvaceae. We demonstrate that the similarity between at least one of the geographical colour phenotypes of T. sidoides and co-flowering Malvaceae is adaptive, since the former obtains more pollination services when growing together with its model than when growing alone. Instead of the convergent evolution pattern attributed to Müllerian mimicry, our data rather suggest an advergent evolution pattern, because only T. sidoides seems to have evolved to be more similar to its malvaceous models.     

Natural hybridization among subspecies of <Emphasis Type="Italic">Turnera sidoides</Emphasis> L. (Passifloraceae) revealed by morphological and genetic evidence

Turnera sidoides is a complex of outcrossing, perennial, rhizomatous herbs that is widely distributed in southern South America. Five subspecies are recognized taxonomically based on morphological features and geographical distribution. In certain regions, the areas of distribution of the subspecies overlap partially. In such contact zones, the extent of reproductive barriers among subspecies is still largely unknown, but morphologically intermediate individuals have been found in the field, indicating that hybridization may actually occur between subspecies. Crossability among subspecies of T. sidoides has been shown by experimental studies with cultivated plants, but the mechanisms involved in natural populations are still unknown. To investigate the mechanisms that underlie gene flow within the T. sidoides complex, in this paper we analyze the morphological and genetic variation, as well as the crossability among taxa in a contact zone between subspecies pinnatifida and sidoides, in southeastern Uruguay. Our results constitute the first evidences of ongoing natural hybridization between subspecies of T. sidoides and suggest that, although hybridization may not have been of significance in the early phase of the species differentiation, reticulate evolution is ongoing enhancing the current morphological and genetic variability of the complex.     

Chloroplast DNA variation in the Turnera sidoides L. complex (Turneraceae): biogeographical implications

Aim To analyse the current geographical structure of chloroplast DNA variation in the Turnera sidoides L. complex in order to establish historical biogeographical hypotheses for the mid‐latitude South American lowlands. During the Quaternary, the climate shifted from tropical humid to cold dry, and the vegetation cover has not been stable. The consequences of these processes on the current distribution of the vegetation of this area have received very little attention. Location The mid‐latitude South American lowlands extend between c. 20 and 40°S and include Uruguay, northern, central and eastern Argentina, southern Brazil, and parts of southern Paraguay and Bolivia. They are surrounded by higher‐elevation systems. Methods Turnera sidoides is a well‐studied polyploid complex of perennial rhizomatous herbs occurring throughout the area of interest. We analysed 321 individuals from 79 populations of the five recognized subspecies. We also included progenies from artificial crosses in order to analyse chloroplast inheritance. After screening sequences for four non‐coding chloroplast DNA regions, the trnL–trnF spacer was selected to characterize the collection. Results Three haplotypes can be easily identified, with each differing from the others in two independent characters. A clear geographic structure is revealed when haplotypes are plotted for the complex as a whole regardless of subspecies and cytotype. Three distinct regions can be identified. Main conclusions We propose three putative refugial areas for the Turnera sidoides complex, which are associated with the orographical systems of the region. Ravines and slopes in the Haedo Cuchilla system in northern Uruguay, the elevations of the western side of the area in Argentina, and the eastern Serranías system in south‐eastern Uruguay may each have served as refugia in which the A, B and C haplotypes became fixed during the drier climatic phases. Biogeographical patterns in the area covered by T. sidoides, particularly east of the Uruguay River, have not previously been analysed from a historical perspective.     

Distribution of style and pollen polygalacturonases among distylous and homostylous Turnera and Piriqueta spp. (Turneraceae)

We explore the distribution of a style and pollen polygalacturonase in a number of distylous and homostylous species of Turnera, and two species of Piriqueta (Turneraceae). We show, using immunoblotting with antibodies made against these proteins, that the style polygalacturonase is specific to styles of short-styled plants of all the six distylous species of Turnera we have investigated. Styles of a somatic homostylous mutant derived from a short-styled plant do not possess the style polygalacturonase. Distylous P. caroliniana did not appear to possess this protein. We show that the pollen polygalacturonase, while associated with the short-styled morph in three species, is polymorphic among short-styled plants of T. krapovickasii, and absent from T. joelii, T. grandiflora and P. caroliniana. These data support a role for the style polygalacturonase in distyly, possibly in the incompatibility system, but cast doubt on any role for the pollen polygalacturonase. In concert with the predictions for the mode of origin, and the response of styles of homostylous species to pollen from long- and short-styled plants, we find that none of the homostylous species possess the style polygalacturonase. The pollen polygalacturonase does occur in some homostylous species, but not in others. It is not clear that the pollen polygalacturonase, however, provides a marker for the mode of origin of homostyly.     

Geographic patterns of morphological variation in Turnera sidoides subsp. pinnatifida (Turneraceae)

With the objective of contributing to understanding the patterns of variation within the Turnera sidoides complex, a detailed evaluation of morphological variation along the range of T. sidoides subsp. pinnatifida was performed. A multivariate analysis based on leaf traits and flower colour data enabled differentiation of five morphotypes. Common-garden experiments demonstrated that the morphological variants have a strong genetic basis. It was also found that the morphotypes are geographically structured along the subspecies range, display different habitat preferences, and occur in regions with different climatic regimes. Although these results are suggestive of adaptive differentiation of T. sidoides subsp. pinnatifida, comparisons between morphological and bioclimatic ordinations showed that the patterns observed cannot be fully explained by current climatic conditions. It is proposed that Miocene–Pleistocene events may explain the origin of the five morphotypes and that current climatic and ecological factors may be contributing to the maintenance of the extent and patterns of morphological differentiation in T. sidoides subsp. pinnatifida.     

Nuclear DNA content in the polyploid complex Turnera ulmifolia (Turnera L., Passifloraceae)

Turnera ulmifolia constitutes a well-studied polyploid complex with allo- and autopolyploid species ranging from 2 to 8x. Flow cytometry was used to determine nuclear DNA content, and to estimate 2C- and 1Cx-values with the aim of analysing the genome size in Turnera in terms of polyploid speciation. The 2C-value and 1Cx-value were evaluated in 12 species of the T. ulmifolia complex. Nuclear DNA content was estimated by flow cytometry of nuclei stained with propidium iodide. The 2C DNA content ranged from 1.38 to 1.83?pg in diploids, from 2.67 to 3.96?pg in tetraploids, from 2.73 to 4.31?pg in hexaploids, and from 3.53 to 5.90?pg in octoploids. The 1Cx-value ranged from 0.44 to 0.99?pg. The Turnera ulmifolia complex showed an increase in total DNA content in the ploidy level, but not in the expected proportion. The general tendency indicated a decrease in the 1Cx-value with increasing chromosome number, with T. grandidentata 4x being an outstanding exception. The 1Cx-values in the allooctoploids T. aurelii and T. cuneiformis differed by 1.6-fold from each other, probably as a result of different evolutionary histories following divergence from the last common ancestor.     

Inheritance of spontaneous mutant homostyles in Turnera subulata x krapovickasii and in autotetraploid T. scabra (Turneraceae)

To explore the genetic architecture of distyly in Turnera spp., we determined the inheritance and compatibility behaviour of two spontaneous homostyled mutants. A long-homostyled mutant shoot arose on an otherwise short-styled plant that was an artificial hybrid (Turnera subulata x T. krapovickasii) between two diploid distylous species. The mutation appears to be an allele, SH, of the distyly locus with the dominance relationships, S>SH>s, where S confers the short-styled phenotype, SH confers homostyly in SHSH and SHs genotypes, and ss genotypes are long-styled. Aberrant segregation ratios were observed among some crosses and might be the result of pollen competition. Compatibility relationships are consistent with the hypothesis that a gene complex determines distyly. Infrequently, revertant short-styled flowers have appeared on cuttings of the T. subulata x T. krapovickasii mutant and on occasion, short-styled progeny have appeared in crosses where none were expected. A second mutant homostyle was discovered in autotetraploid T. scabra. The mutation is inherited as above, however, tetrasomic inheritance occurs at the locus. This homostyled mutant carries two copies of the SH allele and has the duplex genotype SHSHss. Compatibility relationships were as observed above. The occurrence of homostyled mutants is consistent with the hypothesis that a linked gene complex underlies the inheritance of distyly in Turnera but we cannot discount the hypothesis that an allelic series is responsible.     

Phylogeny of Geomydoecus and Thomomydoecus pocket gopher lice (Phthiraptera: Trichodectidae) inferred from cladistic analysis of adult and first instar morphology

The phylogeny for all 122 species and subspecies of chewing lice of the genera Geomydoecus and Thomomydoecus (Phthiraptera: Trichodectidae) hosted by pocket gophers (Rodentia: Geomyidae) is estimated by a cladistic analysis of fifty-eight morphological characters obtained from adults and first instars. The data set has considerable homoplasy, but still contains phylogenetic information. The phylogeny obtained is moderately resolved and, with some notable exceptions, supports the species complexes proposed by Hellenthal and Price over the the last two decades. The subgenera G. (Thaelerius) and T. (Thomomydoecus) are both shown to be monophyletic, but the monophly of subgenus T. (Jamespattonius) could not be confirmed, perhaps due to the lack of first-instar data for one of its component species. The nominate subgenus of Geomydoecus may be monophyletic, but our cladogram was insufficiently resolved to corroborate this. Mapping the pocket gopher hosts onto the phylogeny reveals a consistent pattern of louse clades being restricted to particular genera or subgenera of gophers, but the history of the host-parasite association appears complex and will require considerable effort to resolve.     

Systematics of the olive thrush Turdus olivaceus species complex with reference to the taxonomic status of the endangered Taita thrush T. helleri

During the last 40 years, few species of African birds have undergone more taxonomic revision than the olive thrush Turdus olivaceus. This is due to disagreement on how to partition the striking phenotypic variation among allopatric populations. The current consensus is to recognise one species T. olivaceus , split into three assemblages: (1) the olivaceus group restricted to southern Africa, (2) the swynnertoni group of the Zimbabwean and southern Malawi highlands, and (3) the abyssinicus group of the montane highlands of eastern and central Africa. Mitochondrial DNA sequences from 63 individuals were analysed to investigate the phylogenetic relationships among 16 taxa (species and subspecies) in the olivaceus species complex (plus seven outgroup species), with, particular emphasis on the relationships and taxonomic status of the endangered Taita thrush ( helleri ). Phylogenetic hypotheses generated using parsimony, maximum likelihood, and Bayesian inference identified a number of discrete clades corresponding to recognised subspecies. Northern ( abyssinicus clade) and southern populations ( olivaceus + swynnertoni clade) of olive thrush differ by 9–10% in sequence divergence. Furthermore, all analytical methods suggested that helleri (Taita Hills) and roehli (Usambara and Pare Mountains) are reciprocally monophyletic with respect to mtDNA, and 2.5 to 10.5% divergent from all other forms of olive thrush. Both helleri and roehli are surrounded in adjacent highlands by populations of olive thrush that represent a more recent radiation, suggesting that helleri and roehli may be relict taxa which have been able to maintain their genetic integrity. The results of this study support previous arguments for recognizing the arid/woodland T. smithi as a species distinct from other southern African forest populations of T. olivaceus (including the swynnertoni group). Results further suggest that T. abyssinicus , T. helleri , and T. roehli be accorded species rank.     

Evidence of 2n microspore production in a natural diploid population of Turnera sidoides subsp. carnea and its relevance in the evolution of the T. sidoides (Turneraceae) autopolyploid complex

Turnera sidoides is a complex of distylous perennial rhizomatous herbs that includes five subspecies. Since polyploidy has played a prominent role within this species (x?=?7), ongoing studies in T. sidoides are focused on the understanding of the mechanisms involved in the origin and the establishment of polyploids. Therefore, aiming to contribute to the understanding of the mode of polyploid formation, in this study we investigated the frequency of unreduced microspores in a natural diploid population of T. sidoides subsp. carnea by analyzing the size range of pollen and the constitution of the sporads. The results showed that some of the individuals studied produced 2n and 4n microspores, both in short- and long-styled floral morphs. The analysis performed documents the production of unreduced microspores in T. sidoides subsp. carnea and provides evidence that support the hypothesis of sexual polyploidization as one of the most probable mechanisms involved in the origin of polyploids within this species complex. The role of unreduced pollen in the establishment and persistence of newly formed polyploids in diploid populations of T. sidoides is also discussed.     

Chloroplast DNA and the determination of species status in the Disa tripetaloides complex (Orchidaceae), and its relationships to three species Racemosae, section Disa

Disa cardinalis and three populations within the D. tripetaloides species complex contain variation in their chloroplast DNA (cpDNA) variability. All four taxa possessed unique cpDNAs and sequence divergence values ranged from 0.34 to 1.03%. A phylogeny of these genomes was reconstructed, along with the genomes of three other species, D. racemosa, D. uniflora and D. venosa, all of which are also section Disa and series RAcemosae, to determine the relationship of these closely related species to the D. tripetaloides complex. A phylogeny of the taxa using morphological data was also reconstructed. Outgroup comparison was made with D. sagittalis, a member of section Coryphaea. Although the molecular and morphological data were not completely congruent, both data types revealed D. cardinalis, rather than D. tripetaloides ssp. aurata, to be more closely allied with D. tripetaloides ssp. tripetaloides, suggesting that D. tripetaloides ssp. aurata should be elevated to species rank. Additionally, the high sequence divergence observed between the Natal and Cape populations, coupled with their geographical isolation and alternate flowering seasons, suggests that these two D. tripetaloides ssp. tripetaloides populations may, in fact, be more appropriately ranked as subspecies.     

珍稀蝴蝶的亚种分类问题及保护意义：以喙凤蝶属为例

亚种是种的一个亚单位,由于分类学上的不同定义和主观性而受到质疑。然而,在分类学实践中,所有试图用不同的术语取代亚种或完全放弃亚种的尝试都是不可接受的。亚种同样是生物多样性的重要组成,有一定独特性,为自然进程的一部分,具有保护研究价值。本文基于亚种概念、特征认为亚种分类需参考地理隔离(如异域分布)与表型差异2个原则。我们以珍稀喙凤蝶属Teinopalpus为例,系统收集亚种分类文献,采集该属亚种的地理分布、形态描述与差异等信息,综合分析该属蝶类的亚种分类现状、问题与原因。1843-2007年喙凤蝶属金带喙凤蝶T. imperialis及其姊妹种金斑喙凤蝶T. aureus各自记录了8个亚种。然而,部分亚种记录于同一行政区,如金带喙凤蝶T. i. imperialis, T. i. himalaicus与T. i. behludinii 均在中国四川有记录,而金斑喙凤蝶在中国浙江有2个亚种记录,这显示名称应用的不确定性。考虑到两姊妹种同域分布在亚洲东南部,通过比对亚种正模所在地植被群落(biome)、生态区(ecoregion)的一致性判定地理隔离,发现金带喙凤蝶(覆盖3个植被群落)比金斑喙凤蝶(仅覆盖1个)的种下生态位分化程度更高。而据所处生态区的一致性,建议将金带喙凤蝶亚种分类修订为T. i. imperialis, T. i. himalaicus, T. i. miecoae, T. i. behludinii, T. i. imperatrix(含T. i. bhumipholi), T. i.gillesi和T. i. gerritesi 7个;金斑喙凤蝶亚种分类修订为T. a. aureus(含T. a. wuyiensis, T. a. guangxiensis与T. a. nagaoi), T. a. eminens(含T. a. laotiana), T. a. shinkaii与T. a. hainani 4个。由于可参考标本少,喙凤蝶属亚种确立时可用形态比对信息有限、不完整(如只依据单性的形态比对结果等),易“过度亚种化”。珍稀亚种分类问题(如“过度亚种化”、信息不确定性等)影响其保护措施,因为管理者常需权衡投入成本与成效、明确优先保护区域或对象,并以此做出决策。因此,在更多确定信息未获得前,不建议进行亚种分类。     

Trichome complement of Turnera and Piriqueta (Turneraceae)

The indumentum of Piriqueta and Turnera is made up of nine different types of trichomes, which broadly can be divided into glandular and non-glandular. Taking into account foot shape, head size and pedicle size, five variants of glandular trichomes are recognized: microcrapitate, stipitate-capitate, sessile-capitate and setiform. The non-glandular trichomes can be simple (unicellular or pluricellular-uniseriate), stellate or porrect-stellate. The setiform glandular hairs are present in most species of Piriqueta . Simple unicellular hairs are the most widespread type, frequently being found in combination with other trichomes. Stellate trichomes show a restricted distribution in both genera. Within Piriqueta , section Africana has only simple trichomes, whereas section Piriqueta has also porrect-stellate trichomes; groups of species can be set up according to the presence and type of glandular trichomes. Within Turnera the stipitate-capitate trichomes are exclusive to series Papilliferae ; sessile-capitate trichomes are found in series Microphyllae , Annulares and in some species of Salicifoliae ; clavate trichomes are found only in series Turnera ; setiform glandular hairs are exclusive to T. collotricha , whereas the microcapitate trichomes are widely distributed.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 85–97.     

The conservation genetics of Ash Meadows pupfish populations. I. The Warm Springs pupfish <Emphasis Type="Italic">Cyprinodon nevadensis pectoralis</Emphasis>

The Warm Springs pupfish (Cyprinodon nevadensis pectoralis) inhabits several low flow (<0.15 cfs) springs in an otherwise dry Mojave Desert landscape. Increasing demands on groundwater coupled with predictions of decreased precipitation from global warming make this species a sentinel for biodiversity dependent on springs. Here we examine mitochondrial DNA haplotypes and nuclear microsatellite genotypes for individuals sampled in 1998 and 2007 as a means of inferring the historical demography of the subspecies. Estimates of genetic effective population size from comparison of allele frequencies over time underscores that all spring populations support small populations, typically <100 individuals. Such small population sizes suggest spring populations may have heightened probabilities of extirpation. Despite small population sizes, all springs harbor relatively high levels of genetic variation for both nuclear and mitochondrial DNA. Unexpectedly high levels of variation may be explained by recent declines in population size coupled with rare episodes of gene flow between springs suspected of occurring during flooding events. The inferences gained from the genetic data provide the basis for evaluating current and future restoration plans. The genetic data suggest there is a need to balance the predicted positive effects of restoring hydrological connectivity with the potential negative impacts of providing avenues for the spread of exotic species detrimental to pupfish populations.     

Measure of molecular diversity within the Trypanosoma brucei subspecies Trypanosoma brucei brucei and Trypanosoma brucei gambiense as revealed by genotypic characterization

We have evaluated whether sequence polymorphisms in the rRNA intergenic spacer region can be used to study the relatedness of two subspecies of Trypanosoma brucei. Thirteen T. brucei isolates made up of 6 T. b. brucei and 7 T. b. gambiense were analyzed using restriction fragment length polymorphism (RFLP). By PCR-based restriction mapping of the ITS1-5.8S-ITS2 ribosomal repeat unit, we found a fingerprint pattern that separately identifies each of the two subspecies analyzed, with unique restriction fragments observed in all but 1 of the T. b. gambiense "human" isolates. Interestingly, the restriction profile for a virulent group 2 T. b. gambiense human isolate revealed an unusual RFLP pattern different from the profile of other human isolates. Sequencing data from four representatives of each of the two subspecies indicated that the intergenic spacer region had a conserved ITS-1 and a variable 5.8S with unique transversions, insertions, or deletions. The ITS-2 regions contained a single repeated element at similar positions in all isolates examined, but not in 2 of the human isolates. A unique 4-bp [C(3)A] sequence was found within the 5.8S region of human T. b. gambiense isolates. Phylogenetic analysis of the data suggests that their common ancestor was a nonhuman animal pathogen and that human pathogenicity might have evolved secondarily. Our data show that cryptic species within the T. brucei group can be distinguished by differences in the PCR-RFLP profile of the rDNA repeat.     

中国西南部明纹花鼠三个亚种的分化

明纹花鼠是东南亚地区的特有种,在中国有3亚种（阿萨姆亚种、滇西亚种、滇南亚种）,分布于藏东南、云南西南部和南部。以往主要根据外部形态特征进行分类。为探讨3亚种头骨性状的差异,在测量15个陛状的基础上,运用统计分析软件SPSS 11．0对其可测量性状进行多变量分析。结果及推测如下：①明纹花鼠3个亚种的地位是确定的。②在头骨15个性状中,两两亚种之间具有显著差异的性状：阿萨姆亚种与滇西亚种有8个（枕鼻长、吻宽、颤宽、眶间宽、后头宽、鼻骨宽、下齿列、下颌骨高）;阿萨姆亚种与滇西亚种有7个（颅全长、枕鼻长、颧宽、眶间宽、后头宽、下颌高、下颌骨高）;滇西亚种与滇南亚种有4个（吻宽、后头宽、鼻骨宽、下颌高）。③滇南亚种与滇西亚种之间的亲缘关系近：于两者各自与阿萨姆亚种之间的亲缘关系。