Pollen-cones of Pinus bungeana Zucc. ex Endl. (Pinaceae, Coniferales) : Do they indicate a pseudanthial origin?

Even today, the evolutionary origin of coniferous pollen-cones is still controversial and conflicting theories about the identity of their microsporangiophores exist. Previous studies strongly suggest that the simple pollen-cone structure of some Taxaceae s.l. is most likely derived from a compound structure and each microsporangiophore represents a lateral, however markedly reduced flower. To test if a similar evolutionary pathway remembering the pseudathium concept sensu Wettstein applies also for Pinaceae, normal shaped and abnormal pollen-cones of Pinus bungeana (Pinaceae) are investigated with SEM and paraffin microtome technique. Pinus sylvestris, which is the type species of the genus, is used as another example. The early development of bisporangiate microsporangiophores starts with two distinct primordia. They fuse in basal parts and form a common stalk. The distal parts remain free and each develops a stalk, a distinct phyllom-like scutellum and one abaxial microsporangium. In some bisporangiate microsporangiophores two monosporangiate microsporangiophores are inserted laterally at the common stalk, which forms a distinct terminal apex. Other microsporangiophores bear a subunit in form of a second, however, aborted microsporangiophore in a lateral position at the base of the common stalk. It is suggested that the bisporangiate microsporangiophore is not a staminate leaf in the sense of a microsporophyll, but a dorsiventral synangium consisting at least of two fused microsporangiophores. In this case each microsporangiophore corresponds to a markedly reduced cone (= flower) which however has lost its pherophyll (= subtending leaf). Similar as shown for Torreya the simple pollen-cone structure in Pinaceae is thus derived from a pseudanthial (= compound) origin.publishe

The adaptive value of shoot differentiation in deciduous trees and its evolutionary relevance Valor adaptativo de la diferenciación de brotes en árboles deciduos y su relevancia evolutiva

Pronounced long shoot/short shoot differentiation is typical for deciduous angiosperm trees. It also occurs in a number of gymnosperms and very few evergreen angiosperm trees. The study of 719 angiosperm tree species (602 deciduous and 117 evergreen species) demonstrated that the deciduous condition is nearly always associated with shoot differentiation. Detailed measurements in 38 angiosperms showed that the leaf area of an entire short shoot equals the leaf area of a single long shoot leaf of the same species and individual. In the few cases where the leaf area of the short shoot is slightly larger than that of a single long-shoot leaf, the short shoot leaves shade each other and the projection of the short shoot equals the area of a single long shoot leaf. Calculations of the stem biomass needed to expose a given assimilatory surface show two interesting aspects. First, the stem biomass (dry weight) to expose leaf surface is about 10 times less in short shoots than in long shoots. Second, this biomass in long shoots and short shoots appears to be species independent. Regarding shoot structure efficiency, leaf size and shape do not matter. Some evergreen species resemble in all parameters more to deciduous species than to typical evergreen species. Phytogeographical data as well as morphological data suggest that these atypical evergreen species are derived from deciduous ancestors. As measured parameters differ markedly between all gymnosperms, except Ginkgo, and angiosperms, we suppose that the evolutionary pathway leading to shoot differentiation was different for gymnosperms and angiosperms.<br>En Angiospermas arbóreas deciduas, es común encontrar un alto grado de diferenciación entre brotes largos y brotes cortos. También se presenta esta característica en un número de gimnospermas y en muy pocas angiospermas arbóreas siempreverdes. El estudio de 719 especies de angiospermas arbóreas (602 deciduas y 117 siempreverdes) demostró que la condición decidua está casi siempre asociada a la diferenciación de los brotes. Mediciones detalladas en 38 angiospermas demostraron que la totalidad del área foliar de un brote corto es semejante al área foliar de una hoja de un brote largo de la misma especie y del mismo individuo. En los pocos casos en que el área foliar del brote corto es levemente mayor que el área de una hoja de un brote largo, las hojas del brote corto se sombrean entre sí de manera que el área proyectada del brote corto se asemeja a la de la hoja del brote largo. Cálculos de la biomasa de tallo necesaria para soportar una determinada superficie asimilatoria mostraron dos aspectos interesantes. Primero, que la biomasa de tallo (peso seco) de soporte del área foliar is alrededor de 10 veces menor en brotes cortos que en brotes largos. Segundo, que esta biomasa en brotes largos y brotes cortos parece ser independiente de la especie. En cuanto a la eficiencia estructural de los brotes, el tamano y la forma de las hojas no son relevantes. Algunas especies siempreverdes se asemejan en todos sus parámetros más a las especies deciduas que a especies siempreverdes típicas. Datos fitogeográficos así como datos morfológicos sugieren que estas especies siempreverdes atípicas derivaron de ancestros deciduos. Como todos los parámetros medidos difieren notablemente entre todas las gimnospermas, excepto Ginkgo, y las angiospermas, suponemos que el camino evolutivo que condujo a la diferenciación de brotes fue diferente en gimnospermas y angiospermas

Morphology and anatomy of anomalous cladodes in Sciadopitys verticillata Siebold & Zucc. (Sciadopityaceae)

The needles of Sciadopitys verticillata Siebold & Zucc. (Sciadopityaceae) are inserted in the axils of the reduced scaly leaves and are thus generally regarded as cladodes. The evolutionary pathway leading to these cladodes remained, however, dubious and conflicting interpretations are given. Intermediate morphological structures are known basically from one single paper (Carrière 1868), giving only line drawings of the morphology without any further anatomical details. Attempts to produce such intermediate cladodes in experiments failed, but amongst thousands of checked individuals by chance one was found displaying such intermediate or malformed cladodes. These were analyzed morphologically and anatomically in detail. The results gave new insights into the evolution of these cladodes and might raise new questions about the monophyletic origin of the gymnosperm leaf

Comparative embryology and taxonomic considerations in Eriocaulaceae (Poales)

The genus concept within Eriocaulaceae is widely discussed because it is mainly based on a small number of floral characteristics of unknown functional and evolutionary significance. The aim of the present work is to comparatively study the embryology of representatives of most genera of Eriocaulaceae to identify relevant features that might aid in circumscribing them. In Eriocaulaceae, the uniformity of the embryological characteristics makes it difficult to interpret the existing relationships among the genera. Some embryological characteristics, especially those related to the number of microsporangia, are shown to be unstable and restricted to Paepalanthoideae. The unique pollen morphology may be related to mechanical processes to accommodate the increase in volume after early formation of the pollen wall and may indicate a relationship to an inaperturate ancestor. The placentation, formerly stated to be axile, is more likely to be central due to protusions of the locular base. The shape of the proximal region of the megagametophyte may be responsible for the formation of the antipodal cyst. The seed coat structure is uniform in origin and is a consistent taxonomic characteristic of the family. A review of Eriocaulaceae is necessary, in which additional morphological and anatomical characteristics should be considered and combined. © 2010 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

Morphology and anatomy of male cones of Pseudotaxus chienii (W.C. Cheng) W.C. Cheng (Taxaceae)

Results of the present study indicate that male cones of Pseudotaxus chienii are representing inflorescences with strongly reduced flowers. The results fit quite well with investigations showing that sporangiophores of Taxus and also of Pseudotaxus comply with reduced flowers. The only difference between male cones in Taxus and Pseudotaxus is the absence of pherophylls in Taxus. Furthermore our results complete a transition series beginning with Cephalotaxus going on to Pseudotaxus and ending with Taxus and Torreya. In this progression Pseudotaxus can be regarded as an intermediate link between the inflorescences of Cephalotaxus and the simple, unbranched cones of Taxus. The entire transition series shows that sporophyll-like sporangiophores can be derived by reduction of lateral cones. There is however no sign that a similar process has occurred in other conifer groups

Male cone evolution in conifers

Despite the simple structure of male conifer cones, there is an enormous variability in cone properties observed upon more careful examination. The diversity ranges from simple cones to compound cones. Moreover, cones can be distinguished according to different spatial distributions on the tree. Simple cones are distributed either as solitary cones or as fascicular or clustered aggregations, while compound cones only exhibit fascicular or clustered aggregations. Here, we demonstrate that these different spatial distribution patterns correlate with distinct leaf types and variable branching frequencies. Furthermore, we provide new insights into the evolution of the sporangiophore, particularly in Taxaceae. Two notably important and fast-evolving characters of conifers are the number of sporangia per sporangiophore and the number of sporangiophores per cone. We demonstrate, across many species and types of cones, how these characters are able to adjust according to the optimal amount of pollen

Male Cone Evolution in Conifers : Not All That Simple

Despite the simple structure of male conifer cones, there is an enormous variability in cone properties observed upon more careful examination. The diversity ranges from simple cones to compound cones. Moreover, cones can be distinguished according to different spatial distributions on the tree. Simple cones are distributed either as solitary cones or as fascicular or clustered aggregations, while compound cones only exhibit fascicular or clustered aggregations. Here, we demonstrate that these different spatial distribution patterns correlate with distinct leaf types and variable branching frequencies. Furthermore, we provide new insights into the evolution of the sporangiophore, particularly in Taxaceae. Two notably important and fast-evolving characters of conifers are the number of sporangia per sporangiophore and the number of sporangiophores per cone. We demonstrate, across many species and types of cones, how these characters are able to adjust according to the optimal amount of pollen.publishe

Floral development and vasculature in Eriocaulon (Eriocaulaceae) provide insights into the evolution of Poales.

BACKGROUND AND AIMS: Floral developmental studies are crucial for understanding the evolution of floral structures and sexual systems in angiosperms. Within the monocot order Poales, both subfamilies of Eriocaulaceae have unisexual flowers bearing unusual nectaries. Few previous studies have investigated floral development in subfamily Eriocauloideae, which includes the large, diverse and widespread genus Eriocaulon. To understand floral variation and the evolution of the androecium, gynoecium and floral nectaries of Eriocaulaceae, we analysed floral development and vasculature in Eriocaulon and compared it with that of subfamily Paepalanthoideae and the related family Xyridaceae in a phylogenetic context. METHODS: Thirteen species of Eriocaulon were studied. Developmental analysis was carried out using scanning electron microscopy, and vasculature analysis was carried out using light microscopy. Fresh material was also analysed using scanning electron microscopy with a cryo function. Character evolution was reconstructed over well-resolved phylogenies. KEY RESULTS: Perianth reductions can occur due to delayed development that can also result in loss of the vascular bundles of the median sepals. Nectariferous petal glands cease development and remain vestigial in some species. In staminate flowers, the inner stamens can emerge before the outer ones, and carpels are transformed into nectariferous carpellodes. In pistillate flowers, stamens are reduced to staminodes and the gynoecium has dorsal stigmas. CONCLUSIONS: Floral morphology is highly diverse in Eriocaulon, as a result of fusion, reduction or loss of perianth parts. The nectariferous carpellodes of staminate flowers originated first in the ancestor of Eriocaulaceae; petal glands and nectariferous branches of pistillate flowers originated independently in Eriocaulaceae through transfer of function. We present a hypothesis of floral evolution for the family, illustrating a shift from bisexuality to unisexuality and the evolution of nectaries in a complex monocot family, which can contribute to future studies on reproductive biology and floral evolution in other groups