958 resultados para Pollen exine ultrastructure
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旌节花科是东亚特有科,仅有旌节花属1属。本文研究了该科植物的生活习性、外部形态、花粉形态和外壁超微结构以及花器官发生,并根据多个DNA区段的序列探讨了科的系统位置和属下关系。在此基础上,综合各方面的证据,对旌节花科进行了全面的分类学修订,完善了该科的分类系统。主要内容包括: 1、习性和外部形态学 结合室内标本和野外考察,对旌节花科植物的生活习性、叶、花序、花和果实各个性状的变异式样、变异幅度、变异规律及其相关性进行了深入研究,并讨论了它们的分类学价值。结果表明:叶片形状、叶缘齿、花序下的叶片是否宿存以及花序是否具花序梗等性状在种间变异非常大,单个性状对旌节花属属下分类意义不大;而叶片毛被、脉序和结构,花序长度,花的大小和颜色以及果实大小等性状通常较稳定,可作为旌节花属划分种和变种的重要依据。 2、花粉形态学和外壁超微结构 在透射电镜下研究了旌节花属5个代表种和1个外类群(Crossosoma californicum)的花粉外壁超微结构,并在扫描电镜下研究了外类群Crossosomataceae 1个代表种(Crossosoma californicum)的花粉形态学特征,结合前人的研究,探讨了上述性状的分类学意义。结果表明:旌节花属植物花粉形态的共同特征是:花粉粒球形或近球形,三孔沟;外壁具穴状纹饰,穴较细小;外壁内壁分层明显,容易区分(柳叶旌节花除外),覆盖层光滑,几无附属物,柱状层明显,穿孔较少,不形成半覆盖层。各种植物花粉外壁超微结构的特征在旌节花属两个组中较一致,但其差异可作为探讨属下种的分类和系统学关系的依据。此外,对旌节花科和Crossosomataceae花粉表面纹饰和外壁超微结构的研究结果提示:这两个科与省沽油科和山茶科厚皮香亚科有较近缘的关系,而不支持这两个科的姐妹群关系。 3、花器官发生 首次对旌节花属常绿组和落叶组共2个代表种的花器官发生进行了研究,结果表明:该科植物花为功能性单性花,雌雄异株,花发育过程长达10个月,其中2-3个月是冬季休眠期。在花发育早期,雄花和雌花的发育没有差异,花性别的分化发生在花发育后期,即每年春天花开前后的2-3天,雌花中雌蕊的子房开始膨大,柱头伸长,胚珠多数,能育,而雄蕊逐渐干瘪收缩直至最后花粉不育;此时雄花的雄蕊发育良好,花丝粗壮发达,花药饱满,花粉能育,而雌蕊发育逐渐停滞直至败育。花器官发生方式为向心式,发生顺序为:苞片—外轮萼片—内轮萼片—花瓣—外轮雄蕊—内轮雄蕊—雌蕊。花萼4枚,两轮,交互对生,花瓣和雄蕊均4枚一轮几乎同时发生,花瓣覆瓦状排列。旌节花属常绿组和落叶组植物的花器官发生发育过程没有差别。 4、基于多个DNA区段序列的系统发育分析 首次选取进化速率较快的5个DNA区段:核核糖体ITS区和叶绿体rps16, ndhF,trnL-F及trnS-G-G区段,对旌节花属分别代表常绿组和落叶组的10种2变种及外类群Crossosomataceae和省沽油科3种植物的相关序列进行测定,并将上述序列合并构建系统树。研究结果表明:旌节花科作为一个单系群得到很好的支持。旌节花属落叶组和常绿组的划分在基于ITS、叶绿体4个区段单独分析及联合分析得到的系统树中没有得到支持,表现在该联合分析得到的树的两个分支与该属植物的习性和外部形态特征无关,而与它们的地理分布密切相关:其中一支主要分布在东亚的偏北部,另一支主要分布在东亚的南部地区,常绿和落叶的种类则混合在这两支中。此外,分子序列分析的结果对该属植物的分类、系统关系、起源和该科的系统位置均有重要意义。系统树显示出分布于台湾的S. sigeyosii与分布于日本的早春旌节花聚成稳定的一支,说明前者与后者,而不是与西域旌节花关系密切。叶绿体基因树与核基因树存在冲突,表明该属经历过网状进化。推测变种披针叶旌节花与云南旌节花可能是杂交起源。根据叶绿体rbcL序列得出的系统树显示旌节花科与产北美的Crossosomataceae的姊妹群关系支持率低于50%。基于最大似然树,运用罚分似然法推算出这两个科分化时间为68.25 ± 10.36 MYA,这两者共同与省沽油科近缘。 5、分类学研究 综合上述研究结果,对旌节花属进行了分类学修订。属下不设立组;确认该属植物有6种6变种,其中中国特有种3个,变种3个。5种5变种被作为新异名,新组合3个,其中2个改级新组合。对西域旌节花、云南旌节花和滇缅旌节花等进行了模式指定。本文还给出了分种和变种检索表、各类群的形态学描述、地理分布图、标本引证和讨论等。
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本研究通过我国CDBI、 KUN、PE、SZ等主要标本馆约3, 500份馆藏标本的研究和野外考察相结合,对我国蔷薇属(Rosa L.)芹叶组(Sect. pimpinellifoliae DC. ex Ser.)植物以及相关组的一些种进行了性状特征、形态和微形态的研究,对该组的一些种的形态特征描述进行了补充,同时给出详细的地理和海拔范围分布图。综合花粉以及种子(瘦果)形态的研究结果重新制订了分种检索表,同时,对该组一些形态相近容易混淆的种进行了对比研究,特别对一直存在争议的绢毛复合体(绢毛蔷薇R. sericea Lindl.和峨眉蔷薇R. omeiensis Rolfe)进行了大量宏观形态特征的研究,并用光学显微镜(LM)和扫描电镜(SEM)对二者的花粉及种子形态、微形态进行对比研究和分析,主要研究内容包括: 1. 芹叶组孢粉研究 对芹叶组的10个种及相关的4个组共17个种(18个样品)的植物花粉进行了光镜和扫描电镜观察和比较研究。研究结果表明:蔷薇属植物花粉粒大小为中等偏小,极轴长23.98[21.82(R. graciliflora Rehd. et Wils.)~29.18(R. tsinglingensis Pax. et Hoffm.)] μm,赤道轴长28.65[24.15(R. graciliflora)~34.70(R. davidii Crép.)] μm;花粉属辐射对称等极单花粉,花粉形态赤道面观呈球形到超长球形;极面观为三裂圆形或近圆形,三孔沟,孔缘加厚,具中部突起的桥状盖。花粉外壁纹饰为条纹状,光镜下形态特征相差不大;在电镜下外壁条纹和脊沟内穿孔的形状、大小和频度等特征,常具组至种水平上的可见变异,可作为组至种水平划分的依据。 根据花粉外壁条纹特征及穿孔形状和数目等特征,本研究将这些植物的花粉归为5个类型,并编制了分组检索表。同时,根据条纹状的清晰度,排列方式、条纹形状、穿孔大小及其频度等方面的差异,各有特点,对该组的10个种编制了分种检索表。 2. 芹叶组种子形态研究 应用光学显微镜和扫描电镜对我国蔷薇属芹叶组14个种及相关组5个组共36种植物的种子宏观形态及种皮微形态特征进行了观察研究。结果显示,蔷薇属种子形态多样,形状分别为肾形、卵形或锥形等;种子颜色以淡棕色、褐色以及土黄色为主;种子大小种间相差悬殊,相对体积为(长×宽×厚)36.66(4.79~114.47) mm3。光镜下,种子宏观形态特征具组内一致性,在扫描电镜下种子表面结构特征因种而异,其纹饰以网纹为主,可分为3种类型,即近平滑型、负网纹型和网纹型。研究结果表明,蔷薇属种子表面纹饰与地理分布关系不大,具有组及种内稳定性。其种子形态、大小、表而纹饰类型等特征可作为蔷薇属组及种水平上的分类依据。 结合蔷薇属花粉形态研究结果,得出蔷薇属种皮微形态特征与花粉外壁纹饰特征相吻合,在代表组及种的特征上具相关性的结论。同时根据种子形态、微形态结构特征的组间区别和种间差异编制了分组及芹叶组14个种的分种检索表。 3. 绢毛蔷薇复合体的研究 通过对大量标本的研究、野外观察以及扫描电镜对绢毛蔷薇复合体的花粉形态和种皮表面结构进行研究,通过对小叶、花粉及种子的形态定量分析结果支持Rowley (1959)的观点,将峨眉蔷薇处理为绢毛蔷薇的一个变种。 综上研究结果得出,蔷薇属植物的小叶片数目、花被基数以及花粉及种子形态等性状是较为稳定的,这些特征可很好的作为分类学依据。 The morphology, pollen exine sculpture and seed coat structure of the species of Rosa sect. Pimpinellifoliae and related sections were studied.About 3,500 herbarium specimens at CDBI, KUN, PE, and SZ were examined. Field work in Sichuan and Yunnan were conducted. Revisions of some species were carried out and a new key to species of sect. Pimpinellifoliae was proposed based on morphology, pollen exine sculpture and seed coat structure, Detailed morphological descriptions, geographical distributions and the altitudinal ranges of some taxa are given. The systematics of the species complex, the Rosa sericea complex (R. sericea Lindl. & R. omeiensis Rolfe), was emphasized. This thesis focused on the following three aspects: 1. Pollen morphology of Rosa sect. Pimpinellifoliae The pollen morphology of 18 samples representing 10 species of the Eurasian Rosa sect. Pimpinellifoliae and 7 additional species of related sections was investigated under LM and SEM. The pollen grains are monadic, actinomorphic, equipolar, medium-sized, spheroidal to perprolate in equatorial view, 3-lobed circular or semi-circular in polar view, crassimarginate, pontoperculate, and with striate exine sculpture. The striate sculpture varies among sections and species. The equatorial axis ranges from 17.97 μm (R. sikangensis) to 29.18 μm (R. tsinglingensis) with an average of 23.98 μm in length, while polar axis varies from 24.15 μm (R. gracilifolra) to 34.70 μm (R. davidii) with an average of 28.65 μm in length. The pollens can be divided into five types based on striate sculpture and a key to the sections sampled was proposed accordingly. The pollen morphology of species of sect. Pimpinellifoliae is more homogeneous and different from other sections sampled and did not support the two-series subdivisions in sect. Pimpinellifoliae. A key is also provided based on characers of pollen morphology among species in sect. Pimpinellifoliae. 2. Seed coat structure of Rosa sect. Pimpinellifoliae The seed coat structure of 39 samples representing 14 species of Rosa sect. Pimpinellifoliae and 12 additional species of related sections was investigated under LE and SEM. The seed relative volume (Length × width × thickness) ranges from 4.79 to 114.47 mm3 with an average of 36.66. mm3. The seeds are reniform, ovate or oblong in shape, with orange-brown, light brown or deep brown color. Seed coat sculpture was reticulate or striate-like reticulate. There was no difference in sculpture character of various speices under LM, while three types of seed coat sculpture were identified under SEM and a key to species based on the seed coat sculpture was provided. The three types of seed coat sculpture were nearly smooth, areolate and reticulate. The study of the seed coat sculpture of same species sampled from different populations showed that characters on the seed coat are stable, and thus the size, shape and seed coat sculpture can be used in species level identification. Interestingly, characters in the seed coat sculpture and the pollen morphology in sect. Pimpinellifoliae are consistent at in specific or sectional levels. A key to the 14 species sampled was given based on seed coat sculpture. 3. The study on Rosa sericea complex The Rosa sericea complex contains R. omeiensis and R. sericea. They are morphologically similar to one another and the systematic status of R. omeiensis has been controversial. In this study we examined large numbers of herbarium specimens of R. omeiensis and R. sericea and conducted field observations in the Hengduan Mts.. We also performed SEM study of pollen morphology and seed coat structure of R. omeiensis and R. sericea. We further carried out intensive morphometric study on the leaflet, pollen, and seed morphology. Our results showed that R. omeiensis should be sunk to be a variety of R. sericea, just as Rowley’s treatment in 1959. In conclusion, the features in the number of leaflet and petal, and the morphological character on pollen and seed are relatively stable. Therefore these characters are very useful in taxon delimition.
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Background and Aims Rheum, a highly diversified genus with about 60 species, is mainly confined to the mountainous and desert regions of the Qinghai-Tibetan plateau and adjacent areas. This genus represents a good example of the extensive diversification of the temperate genera in the Qinghai-Tibetan plateau, in which the forces to drive diversification remain unknown. To date, the infrageneric classification of Rheum has been mainly based on morphological characters. However, it may have been subject to convergent evolution under habitat pressure, and the systematic position of some sections are unclear, especially Sect. Globulosa, which has globular inflorescences, and Sect. Nobilia, which has semi-translucent bracts. Recent palynological research has found substantial contradictions between exine patterns and the current classification of Rheum. Two specific objectives of this research were (1) to evaluate possible relationships of some ambiguous sections with a unique morphology, and (2) to examine possible occurrence of the radiative speciation with low genetic divergence across the total genus and the correlation between the extensive diversification time of Rheum and past geographical events, especially the recent large-scale uplifts of the Qinghai-Tibetan Plateau.Methods The chloroplast DNA trnL-F region of 29 individuals representing 26 species of Rheum, belonging to seven out of eight sections, was sequenced and compared. The phylogenetic relationships were further constructed based on the sequences obtained.Key Results Despite the highly diversified morphology, the genetic variation in this DNA fragment is relatively low. The molecular phylogeny is highly inconsistent with gross morphology, pollen exine patterns and traditional classifications, except for identifying all samples of Sect. Palmata, three species of Sect. Spiciformia and a few species of Sect. Rheum as corresponding monophyletic groups. The monotypic Sect. Globulosa showed a tentative position within the clade comprising five species of Sect. Rheum. All of the analyses revealed the paraphyly of R. nobile and R. alexandrae, the only two species of Sect. Nobilia circumscribed by the possession of large bracts. The crude calibration of lineages based on trnL-F sequence differentiation implied an extensive diversification of Rheum within approx. 7 million years.Conclusions Based on these results, it is suggested that the rich geological and ecological diversity caused by the recent large-scale uplifts of the Qinghai-Tibetan Plateau since the late Tertiary, coupled with the oscillating climate of the Quaternary stage, might have promoted rapid speciation in small and isolated populations, as well as allowing the fixation of unique or rare morphological characters in Rheum. Such a rapid radiation, combined with introgressive hybridization and reticulate evolution, may have caused the transfer of cpDNA haplotypes between morphologically dissimilar species, and might account for the inconsistency between morphological classification and molecular phylogeny reported here.
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During the microspore division in Datura innoxia, the mitotic spindle is oriented in planes both perpendicular (PE) and oblique (OB) to the spore wall against which the nucleus is situated. However, irrespective of polarity, the usual type of hemispherical wall is laid down at cytokinesis and isolates the generative cell from the rest of the pollen grain (type A). In PE spores the vegetative nucleus initially occupies a central position in the pollen grain, whereas in OB spores the vegetative nucleus lies at the periphery of the grain close to the generative cell. In anther cultures initiated just before the microspore division is due to take place, no marked change can be observed in either orientation or symmetry of the mitotic spindle when the spores divide. In some, however, cytokinesis is disrupted and deposition of the hemispherical wall arrested. In the absence of a complete wall, differentiation of the generative cell cannot take place and binucleate pollen grains are formed having 2 vegetative-type nuclei (type B). The 2 nuclei in the B pollens are always situated against the pollen-grain wall, suggesting that the disruption phenomenon is related to the OB spores. The incomplete wall always makes contact with the intine on the intine-side of the spindle. Wall material may be represented merely as short stubs projecting out from the intine into the cytoplasm, in which event the 2 nuclei lie close to each other and are separated by only a narrow zone of cytoplasm. In other grains the wall is partially developed between the nuclei and terminates at varying distances from the tonoplast; in these, the nuclei are separated by a wider zone of cytoplasm. The significance of these binucleate grains in pollen embryogenesis is discussed.
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In young pollen grains of Datura innoxia, a wall of the usual hemispherical type separates the 2 gametophytic cells initially and, in the electron microscope, appears as an electron-translucent matrix which is contiguous with the intine. Before detachment of the generative cell from the intine, the matrix decreases in thickness and in places is dispersed altogether leaving the plasmalemmae on either side of it in close apposition. A particularly prominent zone, triangular in profile, is left where the wall joins with the intine. After detachment of the cell, remnants of the matrix can be seen distributed irregularly around the cell and it is supposed that these are partly derived from material in the triangular zone as the cell is drawn away from the intine. The wall residues persist throughout the maturation phase of the pollen and are considered to be either callose resulting from incomplete digestion of the initial wall, or some other polysaccharide material which is unevenly laid down along the wall and concentrated at the junction with the intine. In pollen induced into embryogenesis by anther culture, wall material is also distributed irregularly around the detached cell in a series of discrete zones, but these are more extensive than in vivo, closer together and in many instances highly dilated. The wall profiles thus have a beaded appearance, the 'beads' being connected together by short links of the 2 apposed plasmalemmae. The contents of the swollen zones have a similar electron density to that of the matrix in vivo but also show traces of a fibrillar component. It is postulated that this unusual swelling is a prelude to dispersal of the wall by disruption of the plasmalemmal links and to the establishment of cytoplasmic continuity between the 2 cells. The significance of such binucleate pollen grains in the formation of non-haploid embryos is discussed.
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Ultrastructural features of embryogenic pollen in Datura innoxia are described, just prior to, during, and after completion of the first division of the presumptive vegetative cell. In anther cultures initiated towards the end of the microspore phase and incubated at 28 degrees C in darkness, the spores divide within 24 h and show features consistent with those of dividing spores in vivo. Cytokinesis is also normal in most of the spores and the gametophytic cell-plate curves round the presumptive generative nucleus in the usual highly ordered way. Further differentiation of the 2 gametophytic cells does not take place and the pollen either switches to embryogenesis or degenerates. After 48-72 h, the remaining viable pollen shows the vegetative cell in division. The cell, which has a large vacuole and thin layer of parietal cytoplasm carried over from the microspore, divides consistently in a plane parallel to the microspore division. The dividing wall follows a less-ordered course than the gametophytic wall and usually traverses the vacuole, small portions of which are incorporated into the daughter cell adjacent to the generative cell. The only structural changes in the vegetative cell associated with the change in programme appear to be an increase in electron density of both plastids and mitochondria and deposition of an electron-dense material (possibly lipid) on the tonoplast. The generative cell is attached to the intine when the vegetative cell divides. Ribosomal density increases in the generative cell and exceeds that in the vegetative cell. A thin electron-dense layer also appears in the generative-cell wall. It is concluded that embryogenesis commences as soon as the 2 gametophytic cells are laid down. Gene activity associated with postmitotic synthesis of RNA and protein in the vegetative cell is switched off. The data are discussed in relation to the first division of the embryogenic vegetative cells in Nicotiana tabacum.
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(Viability, production and morphology of pollen grains for different species in the genus Manihot (Euphorbiaceae)). The objective of this work was to characterize the viability, production and morphology of pollen for different species in the genus Manihot. Floral buds from Manihot accessions were collected from two germplasm banks at Embrapa Cassava & Fruits. The viability of the pollen was assessed via colorimetric, in vitro and in vivo assays. The diameter of the pollen grains was determined by measuring the transversal length of the grain. The experimental design was entirely randomized. Studies on pollen ultrastructure were performed via scanning electron microscopy. Pollen viability was high in the colorimetric tests and intermediate in vivo tests; there was no germination in the in vitro tests. The average production for all accessions was 1,253 pollen grains per floral bud. The size of the pollen grains varied from 132 to 163 pm in the wild accessions, and 129 to 146 pm in the cultivated accessions. The pollen grains for all accessions were very large, apolar, spherical as well as inaperturate, with an exine ornamented with pila organized in a Croton pattern. The wild accessions, in general, produced more and larger pollen grains compared with the cultivated accessions.
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Pollen morphology of 40 species of Rheum, belonging to eight sections, was investigated under LM and SEM. Four new exine patterns were found in the species: a) microcchinate-foveolate, b) rugulate, c) verrucate-perforate, and d) verrucaterugulate ornamentation. In addition, two patterns, the Rheum-type pollens with microechinate-perforate and fine-reticulate, as previously described, were also confirmed in the present study. Based on above study the evolution trends of pollen morphology in the taxa involved were discussed phylogenetically as below. As microechinate-perforate exine pattern existed commonly, the pattern is, therefore, regarded as the most primitive among all the six types. The fine-reticulate type was thought as a derivative type, deriving from the basic micro echinate-foveolate-perforate pattern, and followed by the rugulate and verrucate-perforate ornamentation. The verrucate-rugulate ornamentation should be the most advanced. More than one pollen type often exist in most of the sections in Rheum. The pollen morphology of Rheum was strongly correlated with its geographical and ecological distribution. Three medicinally important species R. officinale, R. palmation and R. tanguticum can be palynologically distinguished by their ornamentations.
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The comparative study of the ultrastructure of the midgut epithelium of stingless bee larvae that eat plant protein (pollen) and animal protein (carrion) throughout the larval phase, shows variations in the digestive cells that are only relative to larval aging and not to the type of larval diet. The cells of older larvae present a cytoplasm with empty spaces that result from emptying of lipid and glycogen stocks, and the presence of autophagic vacuoles. These results are discussed in relation to the hypothesis that variations in the digestive tract of insects may be associated with different diets or phylogeny. We conclude that different diets do not determine cell morphology adaptations in the studied species. As the variations in the ultrastructure of the midgut epithelium are the same in all studied species, including the necrophagous species Trigona hypogea, throughout the larval stage, this sequence of changes seems to be due to different physiological state during larval development.
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Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)
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Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)
Resumo:
The morphology of the exine of Late Cretaceous and Tertiary specimens of Tricolpites reticulatus previously documented from Kerguelen, the Antarctic Peninsula, and the Otway Basin of southeastern Australia has been re-examined and compared with the three pollen types identified in the genus Gunnera. An Antarctic specimen of T reticulatus (Maastrichtian) has a uniform reticulum with elongated lumina, similar to that characterising pollen type 3a of Gunnera macrophylla (subgenus Pseudogunnera). Late Cretaceous (Maastrichtian) Australian specimens of T reticulatus differ; specimens from McNamara resemble pollen of subgenera Pseudogunnera and Milligania of type 3a or type 3b, while specimens of T reticulatus from Princes show more rounded and equidimensional lumina and are therefore tentatively attributed to pollen type 2 found in subgenera Gunnera, Misandra and Panke. Kerguelen Island T reticulatus (Miocene) are distinct from Vega Island specimens: a closer resemblance of Kerguelen T reticulatus and pollen type 2 of extant Gunnera is hypothesised. A comparison between specimens of the North American Tricolpites reticulatus/microreticulatus and pollen of Gunnera is also made. The clear similarity of the North American specimens of Tricolpites microreticulatus and pollen of Gunnera in shape and in the exine surface features of pollen suggests that this taxon should not be separated from T reticulatus but should be treated as a synonym of this species. (C) 2004 Elsevier B.V. All rights reserved.
Resumo:
A progressive global increase in the burden of allergic diseases has affected the industrialized world over the last half century and has been reported in the literature. The clinical evidence reveals a general increase in both incidence and prevalence of respiratory diseases, such as allergic rhinitis (common hay fever) and asthma. Such phenomena may be related not only to air pollution and changes in lifestyle, but also to an actual increase in airborne quantities of allergenic pollen. Experimental enhancements of carbon dioxide (CO) have demonstrated changes in pollen amount and allergenicity, but this has rarely been shown in the wider environment. The present analysis of a continental-scale pollen data set reveals an increasing trend in the yearly amount of airborne pollen for many taxa in Europe, which is more pronounced in urban than semi-rural/rural areas. Climate change may contribute to these changes, however increased temperatures do not appear to be a major influencing factor. Instead, we suggest the anthropogenic rise of atmospheric CO levels may be influential.