Bulb tunic anatomy and its taxonomic implication in Allium L. (Amaryllidaceae: Allioideae)

Abstract

Allium is taxonomically a difficult genus with blurred taxonomic borders at all taxonomic ranks. In this research, anatomy and morphology of bulb tunics in 42 species of the genus representing its 16 currently recognized sections and 6 subgenera (Allium, Cepa, Reticulatobulbosa, Amerallium, Polyprason and Melanocrommyum) were investigated. Our results indicated the following characters to be most informative at sectional and subgeneric levels: features of calcium oxalate crystals and subepidermal cell layer of the outermost bulb tunic, type of tracheids, and bulbils presence. Three main types of crystals, i.e. prismatic, sand and druses, were fairly common among the investigated species except for the members of A. sect. Acanthoprason (A. subg. Melanocrommyum) that often lack crystals. The subepidermal layers of outer tunics were mostly composed of hexagonal, rectangular, or elongated cells but A. paradoxum (A. subg. Amerallium, sect. Briseis) showed the pentagonal type and A. longisepalum (A. subg. Amerallium, sect. Molium) the elliptic type of subepidermal cells. The members of A. sub. Melanocrommyum showed various types of tunic cells and crystals.     

Cypsela micromorphology and anatomy in Cirsium sect. Epitrachys (Asteraceae,Carduoideae) and its taxonomic implications

Cirsium Miller, one of the largest genera of the Asteraceae (Carduoideae), contains more than 250 taxa. Turkey is one of its diversity centers and according to recent studies this genus is represented by 80 taxa. In the present study, cypsela morphologies, micromorphologies and anatomical structures of 17 C. sect. Epitrachys taxa distributed in Turkey are characterized in detail by using stereomicroscope, light microscope (LM) and scanning electron microscope (SEM). Cypselas are found to be obovate to narrowly oblong, or rarely oblanceolate. Based on SEM characters, six cypselas surface ornamentation types are identified, pericarp anatomical structures are described, as well as the structure of the testa. Crystal formations were observed in three taxa and secretory materials in the mesocarp were evaluated. Data obtained from this study are compared with previously published results and it is concluded that fruit micromorphological and anatomical characters are useful for the delimitation of taxa in Cirsium and may be useful for supraspecific classification.     

中国西南地区铁角蕨科植物叶表皮微形态及其系统学意义

利用光学显微镜对12种铁角蕨科植物即大盖铁角蕨、齿果铁角蕨、黑柄铁角蕨、半边铁角蕨、胎生铁角蕨、西南铁角蕨、岭南铁角蕨、剑叶铁角蕨、江南铁角蕨、巢蕨、狭基巢蕨和水鳖蕨植物的叶表皮微形态进行观察比较。结果表明:12种铁角蕨科植物的叶表皮细胞多为不规则型,垂周壁为浅波状、波状或深波状;叶上下表皮均无毛。它们的气孔器类型有7种,为极细胞型、腋下细胞型、不等细胞型、无规则四细胞型,无规则细胞型、横列型和辐射状细胞型。不同种间叶表皮微形态特征表现出一定差异,对属、种的划分有一定分类学意义。对巢蕨属与铁角蕨属、水鳖蕨属与铁角蕨属的亲缘关系进行讨论,为铁角蕨科植物的系统分类研究提供新的资料。     

Weight‐length relationships and condition factor trends for thirty‐eight fish species in trawled and untrawled areas off the coast of northern Sicily (central Mediterranean Sea)

This paper reports the weight‐length relationship (WLR) for thirty‐eight fish species from the Gulf of Castellammare (NW Sicily, Mediterranean Sea). A total of 59 870 fish were collected and measured during four seasonal trawl surveys carried out in 2005 at 10 to 200 m depth. The b parameter of the WLRs ranged from 1.79 (Cepola macrophthalma) to 3.56 (Conger conger). Significant differences in both a and b values among seasons were detected for eight species. Moreover, the condition factors (as weight‐at‐length) of Merluccius merluccius and Mullus barbatus were compared in different area/year combinations characterized by the presence or absence of trawling impact. While M. barbatus showed a higher condition factor in trawled areas/years, possibly due to density‐dependent factors linked to the increased biomass in the no‐trawl area, no such link was detected in M. merluccius.     

Karyotype studies in thirty‐two species of Lilium (Liliaceae) from China1

Fifty‐two populations representing thirty‐two taxa of Lilium L. from China are karyologically analyzed. The results showed that all populations have the same basic chromosome number x=12, and all species are diploid except Lilium tigrinum, which is triploid with 2n=36. In addition, one population of L. sulphureum is aneuploid with 2n=23. The karyotype evolution in the genus Lilium is mainly in terms of alterations of the fine structure of the chromosomes, not in ploidy or basic chromosome number variation. When combining previous studies and our results, we consider the major driving forces for evolution of Lilium to be an increase in karyotypic asymmetry, and unequal translocation and formation of secondary constrictions. The existence of intercalary satellites on the two largest pairs of chromosomes is correlated with the geographic distribution pattern of Lilium across the world. The Hengduan Mountains is a modern diversity and differentiation center of Lilium and it also could be the center of origin for this genus. In addition, some taxonomic conclusions were verified on the subgenus level. Among the species investigated, the chromosome number and karyotypes of 13 taxa were documented for the first time.     

Seed morphology of Linaria species from eastern Spain: identification of species and taxonomic implications

Seed morphology of 15 taxa of Linaria from eastern Spain has been studied using stereoscopic and scanning electron microscopy. Four different morphological types are described according to features of the seed-coat surface. The type that includes winged seeded taxa is divided into three subtypes. Seed morphology of each type and subtype is described, compared, illustrated and discussed according to its taxonomic and biological implications. A key to identify the different species or groups of species is also supplied. 2001 The Linnean Society of London     

Chromosome diversity and evolution in Allium (Allioideae,Amaryllidaceae)

Despite extensive literature on the diversity of karyotypes in Allium is available, no attempt to analyse these data together, within a robust phylogenetic framework, has been carried out so far. Thus, we examined patterns and trends in chromosome evolution across the genus. Based on literature survey, karyo-morphometric features for 207 species belonging to 12 subgenera of Allium were obtained. Included in the data-set were basic chromosome number (x), somatic chromosome number (2n), total haploid (monoploid) chromosome length (THL) and three different measures defining karyotype structure: CV_CI, measuring how heterogeneous are centromeres positions in a karyotype, CV_CL and M_CA, quantifying interchromosomal and intrachromosomal karyotype asymmetry, respectively. Trends in karyotype evolution were analysed by phylogenetic regressions and independent contrasts. Mean karyotypes highlighted differences and similarities in karyotype structure between the 12 subgenera. Further differences were noted when the two parameters for analysing karyotype asymmetry were assessed. In addition, by examining the effects of increasing karyotype dimensions (a proxy for genome size) on karyotype structure and asymmetry, it was shown that in Allium species, the DNA was added proportionally to their arm lengths. Overall, p = 8 and somehow intermediate karyotype asymmetry levels seem to represent plesiomorphic character-states in Allium.     

Seed coat micromorphology of South American species of Hybanthus (Violaceae)

The seed‐coat of several species of Hybanthus subg. Ionidium from South America were studied by SEM (scanning electronic microscopy). Three different levels of sculpture were observed on the seed coat. The micro‐morphological patterns showed great variation between the species and should be considered as additional characters in future taxonomic treatments. The presence of a conspicuous elaiosome was observed; this structure is well known in the genus Viola and is most likely related to seed dispersal mediated by ants.     

国产18种鳞盖蕨属植物叶表皮微形态特征及其系统学意义

鳞盖蕨属是一个自然类群,由于各类群之间的形态差异以及种内变异比较大,因此其分类一直存在争议。该研究利用光学显微镜(LM)和扫描电子显微镜(SEM)对鳞盖蕨属18种植物(含3变种)的叶表皮微形态进行了观察和比较,并结合形态学、孢粉学和分子系统学探讨其系统学意义。结果表明:在光学显微镜下,18种鳞盖蕨属植物叶上、下表皮的脉上均有毛,叶表皮细胞都为不规则形,垂周壁为深波状或波状; 气孔都分布在下表皮,气孔器类型主要有极细胞型和腋下细胞型两种,叶表皮微形态特征支持鳞盖蕨属是碗蕨科中的一个单系类群; 叶上、下表皮脉间毛的变异特征支持光叶鳞盖蕨、毛叶边缘鳞盖蕨和二回边缘鳞盖蕨处理为边缘鳞盖蕨变种; 在扫描电镜下,叶上表皮角质层多为脊状凸起,多数还具丝状纹饰,角质膜特征与叶片回数有一定的相关性; 共环极细胞型、聚腋下细胞型、不等细胞型和不规则四细胞型只在少数种出现,气孔外拱盖多数凹陷,光滑或有颗粒; 气孔外拱盖内缘平滑、浅波状或齿状; 保卫细胞两极大多数有“T”型加厚,不同种间叶表皮微形态特征表现出一定差异。该研究结果为其分类、演化、系统位置等进一步研究提供了参考。     

Isolation and characterization of eight microsatellite loci from Phaedranassa tunguraguae (Amaryllidaceae)

Phaedranassa tunguraguae is an endangered species endemic to Ecuador. Eight highly polymorphic microsatellite loci were isolated from an enriched genomic library for this species. Levels of polymorphism were evaluated using a total of 31 individuals from a single natural population. An average of 14.1 alleles per locus was detected, and observed heterozygosity ranged from 0.387 to 0.903. All but one locus depart significantly from Hardy–Weinberg equilibrium. These loci are the first microsatellite primers isolated for Amaryllidaceae and will be utilized to investigate patterns of genetic variation of P. tunguraguae, which will contribute data relevant to the conservation of the species.     

Pollen morphology and taxonomic implications of Muscari Miller (Hyacinthaceae) species from Sicily

Pollen morphology of the native Sicilian species of the genus Muscari, one of which is endemic to Sicily, belonging to the subgenera Muscari, Leopoldia and Pseudomuscari, was studied using a light microscope and a scanning electron microscope. Pollen grains are monads, heteropolar and monosulcate; the exine is semitectate reticulate and perforate. The sulcus extends from distal to proximal face and its length is not directly proportional to the size of the granules and varies from one species to another.     

Genetic diversity of Allium munzii (Amaryllidaceae), a rare southern California species and implication for its conservation

An understanding of genetic diversity within and among populations of rare plant species is a prerequisite to develop effective conservation management strategies and reintroduction programs. Allium munzii is a narrow endemic species distributed in western Riverside County, California, USA and known from 18 extant element occurrences. We sampled 119 individuals from 11 element occurrences and investigated within and among population genetic diversity using two variable chloroplast markers (rpL32–trnL intergenic spacer and rpoC1 intron). Of the total genetic variation detected in A. munzii, 87.65% was due to differences among occurrences. Furthermore, our results revealed that most of the element occurrences are strongly genetically differentiated. There are low levels of gene flow between occurrences, not due to isolation by distance but possibly resulting from habitat fragmentation. Non-significant values of Tajima's D and Fu's Fs were found in all occurrences suggesting no demographic expansion in A. munzii. Ex situ seed and bulb conservation is recommended to enable introduction of individuals to occurrences with low abundance and genetic diversity.     

A new species of the wild dragon tree, Dracaena (Dracaenaceae) from Gran Canaria and its taxonomic and biogeographic implications

The arborescent taxa of Dracaena which form the dragon tree group comprise five species found in Macaronesia, Morocco (D. draco), East Africa (D. ombet, D. schizantha), Arabia (D. serrulata) and the island of Socotra (D. cinnabari). A new species of dragon tree, Dracaena tamaranae A. Marrero, R. S. Almeida & M. Gonzalez-Martin, is described from Gran Canaria, Canary Islands. This new species differs from D. draco, the only other Dracaena species currently known in Macaronesia, in having a growth form and inflorescence type and leaves more similar to the East African and Arabian species of Dracaena. In contrast, D. draco appears to be related to D. cinnabari. In this paper, we also present a study of the taxonomy, habitat and ecology of all the species of the dragon tree group. These are found in thermo-sclerophyllous plant communities of tropical-subtropical regions which are rather xerophilous and have a rainfall range of 200–500 mm. Our study indicates two independent colonization events for Dracaena in Macaronesia. In addition, we suggest that the dragon tree group provides an example of two major biogeographical disjunctions between East and West Africa. We postulate that this group has a Tethyan origin, a hypothesis supported by fossil and palaeoclimatic data, and thus parallels the distribution and dispersal pattern of other taxonomic groups.     

Length–weight relationships of thirty‐seven fish species from the Hongshui River,Southwest China

Length‐weight relationships (LWR) were estimated for 37 fish species belonging to ten families from the middle and lower stream of the Hongshui River, one of the main tributaries to the Pearl River, Southwest China. Length‐weight relationships for 26 species are unknown to FishBase and new maximum lengths are recorded for 20 species. These results are useful for fishery research, conservation and management in the Hongshui River.     

Morphology, anatomy, palynology and nutlet micromorphology of the rediscovered Turkish endemic Salvia ballsiana (Lamiaceae) and their taxonomic implications

Salvia ballsiana (Rech. fil.) Hedge, previously known only from the type gathering, is a local endemic species from Turkey. The species was first collected from Gölbaşı (Malatya) by E. K. Balls in 1935, and was not collected again until 2008, when we found it in Gerger (Adıyaman). The diagnostic morphological characters of this rare endemic species are discussed, including its anatomical, palynological and nutlet micromorphological features. Morphological characteristics of leaves, calyces, corollas and types of stamens are useful for sectional and specific delimitation in Salvia . Anatomical characters such as number of ray rows in roots, distance between vascular bundles in stems, mesophyll structures in leaves, shape of mid-rib and presence/absence of sclerenchymatic tissue in petioles are of taxonomic significance. In addition, size, shape and ornamentation of pollen grains and nutlets are diagnostic.     

Seedling morphology of three mangrove species and its taxonomic implications

Seedling establishment is an important part of the life cycle of mangroves. Very early in the juvenile stage, they face adverse situations such as a variable and unstable substrate as well as a high-saline environment Therefore, the propagules should have some ability to float in saline water. Seedlings differ morphologically from the adult plants. Here, I examined the seedling growth stages in three species of mangrove:Aegialitis rotundifolia Roxb.,Aegiceras corniculatum (L) Blanco, andAvicennia marina (Forsk.) Vierh. that grow in the Sundarbans swamps of West Bengal. In all cases, fruits were the propagating units; seeds were inseparable from those fruits. Germination was rapid and required no dormancy period. The hypocotyi pierced the seed coat rather than emerging from the pericarp — a variation on the traditional viviparous type of germination. Seedlings then became exposed prior to abscission from parent plants. Based on germination patterns and differences in seedling morphology observed here, these species should be classified in a more relevant taxonomic group than was done previously.     

Phylogenetic relationships of Iranian Allium sect. Allium (Amaryllidaceae,Allioideae) as inferred from nrDNA ITS,cpDNA rps16 and trnL–F sequences

Allium is a particularly species rich (more than 800 species) and economically important genus, with numerous taxonomic problems at all levels of classification. In this study, we try to uncover the phylogenetic relationships of the common leek Allium ampeloprasum based on selected samples of this species and its putative relatives in A. sect. Allium from Iran. The silica‐dried leaf samples of 56 accessions representing 23 species of Allium were sequenced; 53 sequences of nrDNA ITS, 35 sequences of plastid rps16 and 52 sequences of trnL–F were generated and additional accessions were extracted from GenBank in order to cover all recognized main lineages in the genus. Maximum Parsimony and Bayesian Inference generated similar trees, but the placement of A. ampeloprasum and its relatives differed slightly between the nuclear and the plastid phylogenies. In the nrITS tree, A. ampeloprasum is retrieved as a highly supported clade with A. iranicum, while in the combined plastid tree A. ampeloprasum formed a highly supported clade with A. vineale. This supports the hypothesis of a possible hybrid origin of A. ampeloprasum. Allium iranicum formed a clade in the plastid tree, but was resolved as paraphyletic in the nrITS tree, probably due to presence of multiple non‐concerted copies of nrITS. Close relationships are suggested between the following species: A. aznavense and A. wendelboi with A. talyschense, A. erubescens and A. rotundum with A. scorodoprasum and A. abbasii with A. phanerantherum.