Systematic significance of tridentate pedicellariae in the echinoid genera Diadema and Echinothrix

Abstract. The tridentate pedicellariae in all species of Diadema and Echinothrix recognized by Mortensen, plus species described later (to include D. palmeri and two color morphs of E. calamaris) were examined to identify diagnostic species characters, and to determine whether such characters are useful in inferring phylogenetic relationships between species. Nineteen morphological characters were measured and analyzed, and species‐specific characters were identified. The morphometric data were then transformed using gap coding and a parsimony analysis was undertaken. The resulting cladogram for the tridentate pedicellariae present in species of Diadema was compared with the mitochondrial DNA phylogeny of Lessios et al., with a good level of congruence observed. Very narrow forceps‐like tridentate pedicellariae were found to be basal among Recent species, present only in D. setosum and D. palmeri. Members of D. palmeri were found to be unique within the genus, having both the forceps‐like form and a broad form of tridentate pedicellaria. The very narrow forceps‐like form is absent in all subsequent divergent species, while the broad form of tridentate pedicellaria in D. palmeri is basal to the broad forms in D. antillarum, D. mexicanum, D. paucispinum, and both the broad and narrow forms of tridentate pedicellaria of D. savignyi. The greatest similarity was found between the broad forms of tridentate pedicellaria in D. antillarum and D. mexicanum. These species were also found to have narrow forms of tridentate pedicellaria that showed a reasonable level of association to one another and to the rostrate form in D. ascensionis. Since the work of Mortensen, D. ascensionis has been shown, using mitochondrial DNA, to be nested within D. antillarum. The results of this study show that the tridentate pedicellaria in D. ascensionis are distinctly different from all other forms within the genus, to the extent that they belong to a rostrate subclass. This indicates that, among Diadema species, the tridentate pedicellariae are one of the first morphological characters to diverge in genetically isolated populations. Unlike the tridentate pedicellariae in Diadema, the different forms of tridentate pedicellariae in E. calamaris and E. diadema showed a stronger association intraspecifically than between forms interspecifically. The small forms of tridentate pedicellaria of the white and brown color morphs in E. calamaris showed a high level of association, while the large forms showed only a moderate association, far lower than would be expected between color morphs of the same species. This suggests that they are either subspecies or recently diverged separate species. This study demonstrates the taxonomic value contained within the characters of the tridentate pedicellariae for both differentiating species (even subspecies) and inferring phylogenetic relationships between species.     

Taxonomic importance of seed morphology in Gentiana (Gentianaceae)

The seeds of Gentiana show high diversity in size, shape and surface pattern. Until now, seeds of only a limited number of taxa have been studied in detail and their utility in the subgeneric classification of Gentiana has not been evaluated. In this study we investigated seed micromorphology of the poorly known sections Pneumonanthe, Frigidae and Isomeria. In order to evaluate the relevance of seed sculpturing for taxonomy, we selected qualitative characters of seed shape and testa ornamentation of all documented sections and performed cluster analyses based on presence and absence of coded characters. Several new scanning electron microscopy pictures of seeds provided a number of valuable characters for the subgeneric differentiation of Gentiana. The results of the cluster analyses of seed characters generally support the sectional classification given by former authors. However, the European Gentiana asclepiadea does not group together with other taxa of section Pneumonanthe, but clusters near to the strictly European section Gentiana. Gentiana froelichii, endemic to the south‐east Alps and treated traditionally within section Frigidae, clusters together with taxa of the European section Ciminalis. Our results reveal that seed micromorphology supports the reassignment of some taxa incertae sedis within Gentiana. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 101–115.     

Pollen morphology within the Monodora clade,a diverse group of five African Annonaceae genera

Pollen morphology has played a major role in elucidating infrafamiliar‐level systematics and evolution within Annonaceae, especially within the African genera. The Monodora clade is composed of five genera, Asteranthe, Hexalobus, Isolona, Monodora and Uvariastrum, which are restricted to Africa and contain together c. 50 species. A molecular phylogeny of the family showed that the monophyly of the Monodora clade is strongly supported and that it is part of a larger clade of 11 African genera. In order to support classification a detailed survey was made of the pollen morphological variation within the Monodora clade, using scanning and transmission electron microsopy. For the two most species‐rich genera, Isolona and Monodora, a molecular species‐level phylogeny was used to assess the taxonomic usefulness of the pollen characters. The survey showed a wide range of pollen morphological diversity. The most conspicuous variation concerned the occurrence of monads without a thicker outer foliation in the basal exine layer in Isolona in contrast to tetrads with a thicker outer foliation in Asteranthe, Hexalobus, Monodora and Uvariastrum. At the infrageneric level, Hexalobus, Isolona and Monodora showed the largest diversity, with various pollen types based on tectum morphology. Hexalobus is exceptional with three types within only five species. The pollen types defined in this study are hardly useful in characterizing major groups identified within both Isolona and Monodora, but they do illustrate relationships within smaller groups.     

Extraocular vision in the sea urchin Diadema setosum

In most organisms, specific structures are responsible for detecting light patterns and vision. Several species of sea urchins appear to have a diffuse photoreceptor system that enables them to detect light and in some cases objects. The presence of extraocular vision was investigated on a sea urchin common in northern Oman, Diadema setosum. Urchins from 8- to 10-m deep coral communities were used in controlled experiments that quantified the orientation response of the urchins to various visual cues: size, contrast and colour of circular targets simulating urchins’ outlines. Urchins responded to black discs down to 11° initial visual angle but not to smaller discs. The minimum grey-scale value triggering a response was between 50 and 37% black. Urchins responded to red targets but not to green or blue ones. An average angular distance between spines of 5.3° suggested a visual acuity of about 10.6°. D. setosum thus showed extraocular vision allowing them to detect objects of different sizes, contrasts and to some extent colour, supporting the hypothesis that the whole sea urchin’s body acts as a single large compound eye.     

淫羊藿属花粉的外壁纹饰及其分类学意义

利用扫描电子显微镜对小檗科淫羊截属,３４种及亚种４４个花粉样品进行了观察和研究。该属植物花粉的外壁纹饰有由不规则条纹状突起到穴状,穴网状,再至网状或条网状学化的趋势,其中包括穴由少到多,由小到大;网脊表面具微刺至平坦的演化方向,与形态性状的演化基本一致,具有重要的分类价值。     

Taxonomic significance of pollen types in the Guyana Highland-centred composite genera of Mutisioideae (Asteraceae)

The Guyana Highland-centred genera, or Stenopadus group, are a complex of species that belong to the tribes Mutisieae and Stifftieae of Asteraceae. The pollen morphologies of 29 species, from 12 of 13 genera of this complex, are described and illustrated using light and scanning electron microscopy. The exine sculpture and structure are highly significant. Four exine types, previously characterized, were distinguished: Gongylolepis , Wunderlichia , Mutisia , and Stenopadus . The characterization of the Stenopadus exine type is enlarged here. These exine types led to the recognition of four well-defined pollen types, whereas the spine length and exine thickness characterized six subtypes. Pollen types circumscribe genera or groups of genera, and some subtypes distinguish species. The pollen morphology within the complex is discussed in relation to the rest of Mutisioideae and other palynologically allied tribes of Cichorioideae. There is little correlation between pollen types and tribes; only the Stenopadus exine type is exclusive to the Stifftieae tribe. The remaining types are shared by the two tribes of the complex. Pollen morphology supports the hypothesis that this group of genera is close to the Gochnatia complex and the Cardueae tribe.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 327–340.     

Endogenous substrate for energy metabolism and ultrastructural correlates in spermatozoa of the sea urchin Diadema setosum

Energy metabolism in spermatozoa of the sea urchin Diadema setosum of the order Diadematoida was examined. The spermatozoa contained not only several kinds of phospholipids and cholesterol, but also triglyceride (TG). Glycogen and glucose were present at extremely low levels. Following dilution of dry sperm and incubation in seawater, the TG content decreased rapidly. Other lipids, however, remained at constant levels, except for a slight increase in the level of free fatty acid. High lipase activity was demonstrated in the spermatozoa. ¹⁴C-labeled fatty acid was oxidized to ¹⁴CO₂. Ultrastructural study also showed that lipid globules were present at the bottom of the midpiece. After incubation in seawater, morphological changes in the lipid globules were observed and some vacuoles appeared. Thus, the results obtained strongly suggest that D. setosum spermatozoa obtain energy through oxidation of fatty acid from TG stored in the lipid globules at the midpieces. © 1995 Wiley-Liss, Inc.     

Taxonomic significance of variation in sexual condition in Pelekium Mitt. and related genera (Thuidiaceae)

Sexual condition has been employed as a key character within the moss family Thuidiaceae to discriminate the dioicous genera Aequatoriella and Thuidiopsis from the monoicous Pelekium. Here, we implemented phylogenetic analyses of the chloroplast trnL-F region and two unicopy nuclear regions to test the monophyly of Pelekium, and hence, the taxonomic significance of the variation in sexual systems. Aequatoriella was deeply nested within Pelekium, suggesting that the sexual condition is not a fixed character state that can be used to circumscribe genera in the Thuidiaceae and, therefore, that Aequatoriella should be sunk. Incongruence among loci rendered the interpretation of the position of Thuidiopsis challenging, and we suggest maintaining it as a separate genus pending further evidence.     

Pollen morphology of the tribes Naucleeae and Hymenodictyeae (Rubiaceae – Cinchonoideae) and its phylogenetic significance

The tribe Naucleeae has recently been recircumscribed on the basis of both morphological and molecular [ rbcL , trnT-F , internal transcribed spacer (ITS)] evidence, and has been found to be the sister group of the tribe Hymenodictyeae Razafim. & B. Bremer. In order to find pollen morphological support for this new classification, the pollen and orbicules of 65 species, representing 23 Naucleeae and the two Hymenodictyeae genera, were investigated by scanning electron and light microscopy. Naucleeae pollen is very small (< 20 µm) to small (20–30 µm) and its shape in equatorial view is suboblate to spheroidal or, more rarely, subprolate. Three compound apertures are present, each comprising a long and narrow ectocolpus, a circular to slightly lolongate mesoporus, and an often H-shaped endoaperture. The sexine ornamentation is perforate, rugulate, or (micro)reticulate, and supratectal elements are always absent. Apart from the variation in sexine ornamentation, the tribe is rather stenopalynous. The pollen of Hymenodictyeae is very similar to that of Naucleeae. The H-shaped endoapertures often observed probably form a synapomorphy for the clade comprising Naucleeae and Hymenodictyeae. Our pollen morphological observations are not in conflict with the widened delimitation of Naucleeae. Unambiguous pollen support for the recent subtribal or generic concepts of Naucleeae could not be found because of a lack of variation of pollen characters within the tribe. Orbicules are invariably present in the ten Naucleeae taxa investigated. They are spheroidal and smooth or irregularly folded.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 329–341.     

Regulation of dendritic spine morphology by an NMDA receptor-associated Rho GTPase-activating protein, p250GAP

The NMDA receptor regulates spine morphological plasticity by modulating Rho GTPases. However, the molecular mechanisms for NMDA receptor-mediated regulation of Rho GTPases remain elusive. In this study, we show that p250GAP, an NMDA receptor-associated RhoGAP, regulates spine morphogenesis by modulating RhoA activity. Knock-down of p250GAP increased spine width and elevated the endogenous RhoA activity in primary hippocampal neurons. The increased spine width by p250GAP knock-down was suppressed by the expression of a dominant-negative form of RhoA. Furthermore, p250GAP is involved in NMDA receptor-mediated RhoA activation. In response to NMDA receptor activation, exogenously expressed green fluorescent protein (GFP)-tagged p250GAP was redistributed. Thus, these data suggest that p250GAP plays an important role in NMDA receptor-mediated regulation of RhoA activity leading to spine morphological plasticity.     

凤仙花属总状花序组的花粉形态及其分类学意义

总状花序组隶属于凤仙花亚属凤仙花属,以具多花的总状花序、侧生萼片2枚、子房5室、果实为线状圆柱形、种子多数、迈卵形、种皮表面具指状突起等特征区别于该亚属的其他类群。前人对凤仙花花粉形态的研究表明凤仙花属的花粉形态对于属下分类及种间界定具有重要的分类学价值。而对于总状花序组来说,有关花粉形态方面的研究少有报道,为了进一步探讨该组花粉S的形态特征及其分类学意义,该研究应用扫描电镜技术对总状花序组的21种植物的花粉进行了观察。结果表明:总状花序组植物的花粉粒均具4条萌发沟,两侧对称,极面观为矩圆形、长矩圆形或近方形;长赤道面观多为椭圆形至矩圆形,短赤道面观为椭圆形;长、短赤道轴比值多在1.4~2.4范围内;表面具网状纹饰,网眼在花粉粒表面均匀分布或赤道面的网眼较极面观的小,网脊边缘光滑或具波状突起,网眼内具不同密度的颗粒状突起。该组的花粉形态与宏观形态性状相关性较小,但花粉极面观形状、长短赤道轴的比值、网眼的分布、网脊边缘形状、网眼内颗粒物密度等特征具有种水平上的稳定性和特异性,对该组植物种间界定具有重要的意义。     

Taxonomic significance of leaf surface morphology in Aloe section Pictae (Xanthorrhoeaceae)

Leaf surface morphology was analysed in 32 species representing the maculate species complex (the poorly resolved section Pictae) in the genus Aloe (Xanthorrhoeaceae). Few comparative morphological data are available for the complex. Leaf surface and stomatal characters observed by scanning electron microscopy show taxonomically significant interspecific variation. Most species are characterized by irregularly outlined, four‐ to six‐sided epidermal cells, the periclinal walls of which are flat and embellished with micropapillae and the anticlinal walls of which are indicated by channels on the leaf surface. The outer stomatal pore is typically sunken or plane and surrounded by four lobes on the leaf surface that may overarch the epistomatal chamber. The guard cells have distinct outer and inner stomatal ledges. Two geographical groups, comprising southern and east African species, are distinguishable by their leaf surface morphology. These characters are diagnostic in A. ellenbeckii, A. prinslooi and A. suffulta and support changes in the delimitation of A. greatheadii, A. macrocarpa and A. swynnertonii. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 418–428.     

Natural growth lines in echinoid ossicles are not reliable indicators of age: a test using Strongylocentrotus droebachiensis

Abstract. Natural growth lines in the ossicles of echinoids have been used to estimate age, calculate growth curves, and infer population age-structure. However, few studies evaluate whether these bands are added annually–a critical assumption of the aging technique. We tested whether the banding pattern is annual in Strongylocentrotus droebachiensis. Sea urchins were tagged with a fluorescent marker, released into tidepools, and collected 1 year later. We quantified the position of the fluorescent mark relative to subsequent growth bands. In 30 individuals ranging in test diameter from 14 to 77 mm, and in a series of ∼2 mm size intervals, we examined 3 interambulacral plates (aboral, ambital, and oral) and a rotula from Aristotle's lantern. Overall, only 7 sea urchins (23%) added a complete band to all 4 ossicles. In 6 sea urchins (20%) at least 1 ossicle added more than 1 complete band. In many sea urchins, especially those >55 mm in diameter, most ossicles added less than 1 band. The banding pattern in ossicles seriously underestimates age in S. droebachiensis and population parameters inferred from these growth lines are biased. Before using the growth-band aging method in other echinoids, it must be demonstrated that 1 band is added annually for all sizes in a population under field conditions.     

中国槲寄生属花粉形态的研究

首次系统地报道中国槲寄生属10种植物花粉形态。通过光学显微镜和扫描电镜观察,国产槲寄生属种类可按花粉的外壁纹饰分为两大类群,与形态上划分相一致。类群Ⅰ(8种)具不显著或近光滑的外壁纹饰,这一类型又可分为2个亚型:纹饰在花粉各个部位基本上是一致的和纹饰在赤道和极、沟两侧不一致的;类群Ⅱ(2种)外壁具显著的杆状纹饰。但花粉性状对形态上相近的种类鉴定意义不大。还讨论了槲寄生属花粉的可能的演化趋势。     

中国桑寄生科的孢粉学研究

系统报道了中国桑寄生科Loranthaceae33种5变种植物的花粉形态,并与澳大利亚 2属6种植物的花粉形态做了比较。通过光学显微镜和扫描电镜观察,国产桑寄生科花粉外壁 纹饰可明显分为两个类型:一种类型为刺状或条状纹饰,另一种为颗粒状纹饰,这与该科的鞘 花族和桑寄生族两个族相吻合。在鞘花族类型中,3合沟、钝刺状或条状纹饰的花粉是基本类 型,合半沟或孔沟形,刺状纹饰的花粉是较进化的类型;在桑寄生族类型中,等极、3合沟、 颗粒状纹饰的花粉是基本类型,异极、副合半沟-合半沟、3沟形和沟形-短沟形或沟孔形、粗 糙或模糊颗粒状纹饰的花粉是较进化类型。根据萌发孔和纹饰可将桑寄生族类型花粉分为3个 类群:类群I包括五蕊寄生属Dendrophtho、梨果寄生属Scurrula、钝果寄生属Taxillus和大苞 寄生属Tolypanthus;类群II仅包括离瓣寄生属Helixanthera;类群III也仅1属,桑寄生属Lor anthus。在这3个类群中,类群I属于基本的类型,属间花粉差别较小,其中梨果寄生属和钝 果寄生属花粉差别最小,显示出较近的亲缘关系;类群II和类群III皆是较进化类型。     

Fruit and seed morphology of the tribe Orobancheae (Orobanchaceae) genera in Turkey and its taxonomic significance

Seed and fruit morphology of four genera of Orobanchaceae: Cistanche, Diphelypaea, Orobanche and Phelipanche, native to Turkey, were studied using light and scanning electron microscopy. In this study, 100 accessions belonging to 39 species were investigated. The examined fruits had loculicidal capsules, generally dehiscing with 2 slits, and the seeds were small and with reticulate ornamentation. The basic shape of the seeds in most of the studied taxa was oblong, ellipsoid to ovoid, globose or subglobose. The results indicate that characters such as seed size, shape and color are useless for identification of the taxa at the species level. However, the anticlinal wall pattern and periclinal wall ornamentation proved useful for identifying the different genera. Four morphological types could be distinguished: smooth, perforate, microfibrillar and granulate. Type I corresponds to the genus Cistanche and Orobanche anatolica, whereas types II, III and IV were found in Diphelypaea, Orobanche and Phelipanche, respectively. The utility of seed characteristics is discussed in terms of inter‐ and intra‐generic taxonomy.     

Links between morphology and function of the pollen wall: an experimental approach

The wall of pollen grains exhibits morphological variation in many features including apertures, ornamentation and thickness, but the function of these characters remains to be clarified. It has been suggested that they are involved in the accommodation of volume changes (harmomegathy). To investigate this further, we developed a protocol that induces a controlled hydration of the pollen without affecting its metabolism and we applied it to six species differing in their pollen wall morphology. The entry of water caused pollen swelling and volume increase leading to breakage of the wall and/or of the plasma membrane, such that the per cent of intact grains was negatively correlated with the level of hydration. Qualitative and quantitative differences were observed between the species. Breakage of the exine was observed only in pollen lacking apertures and with thin exine. Variation in the exine ornamentation and thickness could explain the interspecific differences observed for the rates of breakage of the plasma membrane. Our results suggest that pollen wall morphology matters for survival and maintenance of pollen integrity further to volume increase due to hydration. We propose a rationale for future studies that should allow disentangling the contribution of different pollen morphological and physiological features to harmomegathy.     

Pollen morphology of some related genera of Vernonieae (Asteraceae) and its taxonomic significance

Pollen morphology is an important source of information to increase systematic resolution in Asteraceae, especially in Vernonieae. Aiming to investigate if palynological traits give support to Caatinganthus, Strophopappus and Xiphochaeta as separate genera from Stilpnopappus, we used cluster analysis followed by a test of group sharpness. Further, ordination analysis was applied to detect informative pollen traits associated with the revealed groups. The analyses evidenced five groups: (G1) Caatinganthus rubropappus as a single-species group; (G2) species of Stilpnopappus; (G3) Xiphochaeta aquatica as a single-species group; (G4) Strophopappus bicolor, S. glomeratus, S. villosus, S. ferrugineus, S. pohlii and S. speciosus; (G5) Strophopappus bullatus and S. regnelli. The interruption in the middle of the muri in apertural lacunae explains the single-species group Caatinganthus rubropappus. The thickness of sexine, the type of apertures (porate or colporate), and spine dimensions (length, thickness and distance from each other) are the traits explaining differences between species of Stilpnopappus and Strophopappus. Equatorial lacunae give support to Xiphochaeta aquatica as a single-species group, despite the smaller size of pollen grains of this species as compared to the others species analyzed. The differences among pollen morphology give support to Caatinganthus, Stilpnopappus, Strophopappus and Xiphochaeta as genera according to the taxonomic classification currently accepted. The used approach was efficient to reveal individual pollen traits informative to explain the sharp groups, and was an effective alternative to the use of “pollen types”.