Leaf evolution in early-diverging ferns: insights from a new fern-like plant from the Late Devonian of China

Background and Aims With the exception of angiosperms, the main euphyllophyte lineages (i.e. ferns sensu lato, progymnosperms and gymnosperms) had evolved laminate leaves by the Late Devonian. The evolution of laminate leaves, however, remains unclear for early-diverging ferns, largely represented by fern-like plants. This study presents a novel fern-like taxon with pinnules, which provides new insights into the early evolution of laminate leaves in early-diverging ferns.Methods Macrofossil specimens were collected from the Upper Devonian (Famennian) Wutong Formation of Anhui and Jiangsu Provinces, South China. A standard degagement technique was employed to uncover compressed plant portions within the rock matrix.Key Results A new fern-like taxon, Shougangia bella gen. et sp. nov., is described and represents an early-diverging fern with highly derived features. It has a partially creeping stem with adventitious roots only on one side, upright primary and secondary branches arranged in helices, tertiary branches borne alternately or (sub)oppositely, laminate and usually lobed leaves with divergent veins, and complex fertile organs terminating tertiary branches and possessing multiple divisions and numerous terminal sporangia.Conclusions Shougangia bella provides unequivocal fossil evidence for laminate leaves in early-diverging ferns. It suggests that fern-like plants, along with other euphyllophyte lineages, had independently evolved megaphylls by the Late Devonian, possibly in response to a significant decline in atmospheric CO₂ concentration. Among fern-like plants, planate ultimate appendages are homologous with laminate pinnules, and in the evolution of megaphylls, fertile organs tend to become complex.     

A new Neocalamites (Sphenophyta) with prickles and attached cones from the Upper Triassic of China

Remains of the extinct sphenophyte (horsetail) Neocalamites are most widespread in the Middle–Upper Triassic and are typically represented by stem and leaf fragments. Here we report on spectacular new finds of Neocalamites from the Late Triassic Yangcaogou Formation in Liaoning Province, China that include bedding surfaces dominated by nearly complete aerial stems with attached leaf whorls and rare bractless cones. They reveal a monopodial growth habit for the stems, which are covered with downward projecting prickles that probably provided protection against herbivores. These features provide the basis for a new proposed species, Neocalamites horridus. The nodes bear whorls of very long leaves mainly free to their bases, and one specimen bears an attached cone on a long peduncle. Identical dispersed cones have also been recovered. The leaves of adjacent monopodial stems most likely interlocked to support growth in large stands akin to the role now played by branches in large modern Equisetum species. The new Chinese Neocalamites is among the most confidently reconstructed species, and indicates a greater diversity of sphenophyte morphology during the Mesozoic than previously realized.     

中国二叠纪煤核中具解剖构造的鳞木类植物:似封印叶属(英文)

报道了产于中国二叠纪煤核中的 3种具解剖构造的鳞木类叶 ,它们都具双木质部束 ,与似封印叶属 (Sigil lariopsisScott)特征一致。与该属已有种进行了对比 ,确认它们为 3个新种 :产于山西太原西山煤田太原组上部 7号煤层 (早二叠世早期 )煤核中的山西似封印叶 (Sigillariopsisshanxiensissp .nov .)和太原似封印叶 (S .taiyuanensissp .nov .)以及产于贵州水城矿区汪家寨组 1号煤层 (晚二叠世晚期 )煤核中的贵州似封印叶 (Sigillariopsisguizhouensissp .nov .)。根据煤核中共生的鳞木类植物其他器官以及欧美植物区鳞木类植物的研究资料推断 ,它们可能属于封印木属 (SigillariaBrongniart)的叶。在国外 (主要是欧美植物区 ) ,封印木属植物主要分布于石炭纪 ,见于二叠纪的封印木很少。华夏植物区的封印木属过去很少发现 ,其叶和生殖器官均未报道过。本文是首次报道华夏植物区具解剖构造的封印木属的叶 ,它们的发现不仅丰富了华夏植物区封印木属植物的内容 ,而且对于研究封印木属的演化以及华夏植物区与欧美植物区鳞木类植物之间的关系也具有重要的意义。     

中国二叠纪煤核中具解剖构造的鳞木类植物:似封印叶属

报道了产于中国二叠纪煤核中的3种具解剖构造的鳞木类叶,它们都具双木质部束,与似封印叶属(Sigillariopsis Scott)特征一致.与该属已有种进行了对比,确认它们为3个新种:产于山西太原西山煤田太原组上部7号煤层(早二叠世早期)煤核中的山西似封印叶(Sigillariopsis shanxiensis sp. nov.)和太原似封印叶(S. taiyuanensis sp.nov.)以及产于贵州水城矿区汪家寨组1号煤层(晚二叠世晚期)煤核中的贵州似封印叶(Sigillariopsis guizhouensis sp.nov.).根据煤核中共生的鳞木类植物其他器官以及欧美植物区鳞木类植物的研究资料推断,它们可能属于封印木属(Sigillaria Brongniart)的叶.在国外(主要是欧美植物区),封印木属植物主要分布于石炭纪,见于二叠纪的封印木很少.华夏植物区的封印木属过去很少发现,其叶和生殖器官均未报道过.本文是首次报道华夏植物区具解剖构造的封印木属的叶,它们的发现不仅丰富了华夏植物区封印木属植物的内容,而且对于研究封印木属的演化以及华夏植物区与欧美植物区鳞木类植物之间的关系也具有重要的意义.     

The Upper Permian articulate Phyllotheca australis from South Africa

RAYNER, R. J., 1992. The Upper Permian articulate Phyllotheca australis from South Africa. The taxonomic position of two fossil articulates, which form an important element of the Permian Gondwana flora, is examined in the light of recent collections from South Africa. Vegetative and fertile axes, dispersed and attached sporangia, in-situ spores, isolated and attached whorls of leaves, rhizomes bearing roots and branches, naked axes and a permineralized axis are all assigned to Phyllotheca australis. The large numbers of axes with attached sporangia allow a reinterpretation of the fertile structure and the specific diagnosis is emended. Although it is recognized that the fossils assigned to this taxon may represent a species complex, there exist no clear morphological discontinuities between the material previously assigned to P. australis and P. indica from widely separated Gondwanan localities. Phyllotheca australis was common throughout Gondwana, and is a reliable index fossil of the Upper Permian.     

New evidence of the reproductive organs of Glossopteris based on permineralized fossils from Queensland,Australia. II: pollen-bearing organ Ediea gen. nov

Ediea homevalensis H. Nishida, Kudo, Pigg & Rigby gen. et sp. nov. is proposed for permineralized pollen-bearing structures from the Late Permian Homevale Station locality of the Bowen Basin, Queensland, Australia. The taxon represents unisexual fertile shoots bearing helically arranged leaves on a central axis. The more apical leaves are fertile microsporophylls bearing a pair of multi-branched stalks on their adaxial surfaces that each supports a cluster of terminally borne pollen sacs. Proximal to the fertile leaves there are several rows of sterile scale-like leaves. The pollen sacs (microsporangia) have thickened and dark, striate walls that are typical of the Arberiella type found in most pollen organs presumed to be of glossopterid affinity. An examination of pollen organs at several developmental stages, including those containing in situ pollen of the Protohaploxypinus type, provides the basis for a detailed analysis of these types of structures, which bear similarities to both compression/impression Eretmonia-type glossopterid microsporangiate organs and permineralized Eretmonia macloughlinii from Antarctica. These fossils demonstrate that at least some Late Permian pollen organs were simple microsporophyll-bearing shoot systems and not borne directly on Glossopteris leaves.     

Leafy branches of Gangamopteris from the Gzhelian–Asselian of westernmost Gondwana

The first reconstructions of glossopterids interpreted this Gondwanan group as arborescent, deciduous plants, with leaves and reproductive structures inserted on short shoots, which were arranged on long branches. The leaves are its most abundant organ in the fossil record, but they are mainly found isolated. The arrangement of the leaves as terminal whorls or tight spirals has been the most accepted phyllotaxis hypothesis. The few examples of leaf impressions preserved in connection with axes correspond mainly to leaves of Glossopteris Brongniart, and mostly without clear evidence of the type of insertion. Several specimens of Gangamopteris McCoy leaves attached to axes from the Bajo de Véliz Formation (Latest Carboniferous–Earliest Cisuralian) facilitate reconstruction of the foliar arrangement of the genus, to date known mostly from isolated leaves. The available evidence from the new specimens confirms a variation from well-spaced to dense helical insertion of the leaves without forming true whorls, and discards the early notion that they were mainly clustered apically on short shoots in a similar manner to the extant Ginkgoales.     

Floral development in Anemoneae (Ranunculaceae)

The floral development of two Clematis species and four Anemone species (including Pulsatilla) (Anemoneae, Ranunculaceae) is described. Shared features are: (1) sepals shortly after initiation broad, crescent‐shaped, as opposed to the other organs, which are narrow and hemispherical; (2) outermost organs of the androecium often smaller than the others and sometimes sterile; (3) carpels ascidiate, with distinctive stalk, stigma papillate, decurrent; the carpels have one median fertile ovule and a few lateral sterile ovules in all species studied; the fertile ovule appears before the carpel closes. Generic differences are: (1) In Clematis, four sepals are initiated in two pairs; sometimes one of the sepals in the second pair appears to be divided into two organs (double position) resulting in a pentamerous perianth; the first eight stamens are positioned in two alternating whorls, the outer whorl alternating with the four sepals. In Anemone, the perianth organs, if five, are initiated in spiral sequence; in the Pulsatilla group of Anemone, six sepals are initiated in two whorls; the first three organs of the androecium (staminodes) alternate with the inner sepals. (2) Further androecial organs are mostly in complex whorls (i.e. including double positions) in Clematis, but in an irregular spiral or in irregular complex whorls in Anemone. (3) Anther maturation is largely centripetal in Clematis, but centrifugal or bidirectional in Anemone. In Clematis macropetala, the outermost organs of the androecium lack anthers and the filaments expand and become petal‐like. In contrast, in the Pulsatilla group of Anemone, these organs retain sterile anthers and become small, capitate organs. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 77–100.     

Periodicity of reproductive growth in lycopsids: An example from the Upper Devonian of Zhejiang Province,China

Monilistrobus Wang et Berry is a fossil lycopsid reported from the Upper Devonian Wutong Formation of Jiangsu Province, China, and includes only the type species Monilistrobus yixingensis Wang et Berry. Along a single fertile axis, Monilistrobus has cone-like structures separated by and alternated with a length of vegetative region, which is an important character here termed as periodicity of reproductive growth. The alternation of fertile and vegetative growth occurs regularly and produces repeated intercalary (non-terminal) fertile zones. New specimens, identified as Monilistrobus cf. yixingensis, from the same formation but of Zhejiang Province extend the geographic distribution of this genus. These new specimens demonstrate morphological variations within this genus, such as the lack of spiny appendages on the leaves, different measurements of leaf bases, and different shapes and positions of cone-like structures. The distribution of intercalary fertile zones, as summarized for the first time in this paper, occurs in the Drepanophycales, the Protolepidodendrales, the Isoëtales, the Lycopodiales, the Selaginellales, and Wuxia Berry, Wang et Cai and Monilistrobus of uncertain affinities. Thus, this peculiar trait might have evolved in the very early evolutionary stage of the lycopsids, as indicated in the basal group Drepanophycales. However, the presence of repeated, two or more intercalary fertile zones along a single fertile axis is demonstrated only in Monilistrobus and the extant Huperzia Bernhardi.     

A new plant with novel leaves from the Upper Devonian of Zhejiang Province,China

The evolution of vascular plants during the Devonian Period has had great impacts on terrestrial ecosystems through innovations of key characters such as leaves, heterospory, seed reproduction, and woods. Here we report a new plant, Qianshouia mira n. gen. n. sp., from the Upper Devonian Wutong (Wutung) Formation of Fanwan section, Changxing County, Zhejiang Province, China. This plant has slender axes which seldom branch. Its leaves are arranged in pseudowhorls, departing from axes with an acute angle. The leaves are strap-shaped and each can be subdivided into proximal, middle and distal portions. At the proximal portion, a middle ridge develops, and at the middle portion this ridge divides at least three times forming four to six daughter ridges, and in between are grooves. The distal portion of leaves (leaf apex) is suggested to be adaxially curved, with no ridges but with forked tips. Qianshouia probably represents a small plant with a shrubby or herbaceous habit. Due to the unique leaf morphology and the lack of fertile structures and anatomy, the exact phylogenetic position of Qianshouia is uncertain. Qianshouia might be a lycopsid in light of the pseudowhorled phyllotaxis, or alternatively, could be a sphenopsid if the dichotomous ridge system of the leaf could be demonstrated to be leaf veins. Nevertheless, Qianshouia adds to the diversity of leaf morphologies among the Late Devonian vascular plants.     

Genetic interactions among floral homeotic genes of Arabidopsis.

We describe allelic series for three loci, mutations in which result in homeotic conversions in two adjacent whorls in the Arabidopsis thaliana flower. Both the structure of the mature flower and its development from the initial primordium are described by scanning electron microscopy. New mutations at the APETALA2 locus, ap2-2, ap2-8 and ap2-9, cause homeotic conversions in the outer two whorls: sepals to carpels (or leaves) and petals to stamens. Two new mutations of PISTILLATA, pi-2 and pi-3, cause second and third whorl organs to differentiate incorrectly. Homeotic conversions are petals to sepals and stamens to carpels, a pattern similar to that previously described for the apetala3-1 mutation. The AGAMOUS mutations, ag-2 and ag-3, affect the third and fourth whorls and cause petals to develop instead of stamens and another flower to arise in place of the gynoecium. In addition to homeotic changes, mutations at the APETALA2, APETALA3 and PISTILLATA loci may lead to reduced numbers of organs, or even their absence, in specific whorls. The bud and flower phenotypes of doubly and triply mutant strains, constructed with these and previously described alleles, are also described. Based on these results, a model is proposed that suggests that the products of these homeotic genes are each active in fields occupying two adjacent whorls, AP2 in the two outer whorls, PI and AP3 in whorls two and three, and AG in the two inner whorls. In combination, therefore, the gene products in these three concentric, overlapping fields specify the four types of organs in the wild-type flower. Further, the phenotypes of multiple mutant lines indicate that the wild-type products of the AGAMOUS and APETALA2 genes interact antagonistically. AP2 seems to keep the AG gene inactive in the two outer whorls while the converse is likely in the two inner whorls. This field model successfully predicts the phenotypes of all the singly, doubly and triply mutant flowers described.     

Heterotopic expression of class B floral homeotic genes supports a modified ABC model for tulip (Tulipa gesneriana)

In higher eudicotyledonous angiosperms the floral organs are typically arranged in four different whorls, containing sepals, petals, stamens and carpels. According to the ABC model, the identity of these organs is specified by floral homeotic genes of class A, A+B, B+C and C, respectively. In contrast to the sepal and petal whorls of eudicots, the perianths of many plants from the Liliaceae family have two outer whorls of almost identical petaloid organs, called tepals. To explain the Liliaceae flower morphology, van Tunen et al. (1993) proposed a modified ABC model, exemplified with tulip. According to this model, class B genes are not only expressed in whorls 2 and 3, but also in whorl 1. Thus the organs of both whorls 1 and 2 express class A plus class B genes and, therefore, get the same petaloid identity. To test this modified ABC model we have cloned and characterized putative class B genes from tulip. Two DEF- and one GLO-like gene were identified, named TGDEFA, TGDEFB and TGGLO. Northern hybridization analysis showed that all of these genes are expressed in whorls 1, 2 and 3 (outer and inner tepals and stamens), thus corroborating the modified ABC model. In addition, these experiments demonstrated that TGGLO is also weakly expressed in carpels, leaves, stems and bracts. Gel retardation assays revealed that TGGLO alone binds to DNA as a homodimer. In contrast, TGDEFA and TGDEFB cannot homodimerize, but make heterodimers with PI. Homodimerization of GLO-like protein has also been reported for lily, suggesting that this phenomenon is conserved within Liliaceae plants or even monocot species.these authors contributed equally to this work     

Early Oligocene Callitris and Fitzroya (Cupressaceae) from Tasmania

This paper documents Early Oligocene fossilized foliage and ovulate cones from Lea River, Tasmania and identifies them as belonging to two extant southern hemisphere Cupressaceae genera, Callitris and Fitzroya. Most importantly, it sheds some light on evolutionary trends within Callitris, a genus with numerous extant Australian species and two extant New Caledonian species. Callitris has a very poor fossil record and, because of the present absence of a molecular data set that includes all species, its phylogeny remains somewhat ambiguous. Although Fitzroya foliage has previously been described from a number of Tasmanian sites, this is the first recording of fertile material. The ovulate cones of the Callitris and Fitzroya macrofossils are characterized by bract-scale complexes in two whorls of three and are subtended by scale-like leaves in whorls of three. The fossilized foliage specimens consist of scale-like leaves in whorls of three. These morphological characteristics are only exhibited by three extant southern hemisphere cupressaceous genera, Callitris, Actinostrobus, and Fitzroya. The assignment of the fossils to extinct Callitris and Fitzroya species is made by comparisons with species from these three extant genera. Although much of the Lea River flora are wet rainforest taxa, the Callitris fossils have characteristics of both wet- and dry-adapted extant species.     

Pollen Organ Telangiopsis sp. of Late Devonian Seed Plant and Associated Vegetative Frond

Pollen organ Telangiopsis sp., associated with but not attached to vegetative fronds, has been collected from the Upper Devonian (Famennian) Wutong Formation, Dongzhi County, Anhui Province, China. Fertile axes with terminal pollen organs are dichotomous for 2–4 times and may be proximally attached by fragmentary pinnules. Pollen organs are synangiate and borne on the top of a short stalk. Synangia are radial in symmetry and each consists of 4–8 elongate microsporangia fused at base. Microsporangia have a longitudinal dehiscence line and show a tapered apex. The associated stem is spiny and bears a vegetative frond which bifurcates once at the basalmost part. Frond rachises possess one order of pinna arranged alternately. Pinnules are borne alternately, planate, highly dissected, and equally dichotomous for 2–3 times. Comparisons among Late Devonian seed plants recognize several branching patterns in the fertile fronds/axes bearing terminal pollen organs. Telangiopsis sp. reinforces that the Late Devonian pollen organs are synangiate usually with basally fused microsporangia. It is suggested that the evolutionary divergence of radial and bilateral symmetries of pollen organs may have occurred in the Famennian, when the earliest seed plants evolved planate and sometimes laminate pinnules.     

Merosity in flowers: Definition,origin, and taxonomic significance

The term merosity stands for the number of parts within whorls of floral organs, leaves, or stems. Trimery is considered to be a basic condition that arose through the cyclisation of a spiral flower. Pentamery is mostly derived from trimery by the repetitive fusion of two different whorls. Dimery is either directly derived from trimery, or through pentamery as an intermediate stage. Tetramery is linked with pentamery and should not be confused with dimery. Possible causes for a change in merosity are the reduction of the number of carpels and zygomorphy in flowers. Derivations of different merosities have important consequences for the arrangement of the androecium (the insertion of stamen whorls, their identifications, and their number). It is concluded that two main groups can be identified within the angiosperms: magnolialean and monocotyledonean taxa are mostly trimerous or dimerous; non-magnolialean dicots are mostly pentamerous or tetramerous.     

Male and female flowers of the dioecious plant sorrel show different patterns of MADS box gene expression.

Male and female flowers of the dioecious plant sorrel (Rumex acetosa) each produce three whorls of developed floral organs: two similar whorls of three perianth segments and either six stamens (in the male) or a gynoecium consisting of a fertile carpel and two sterile carpels (in the female). In the developing male flower, there is no significant proliferation of cells in the center of the flower, in the position normally occupied by the carpels of a hermaphrodite plant. In the female flower, small stamen primordia are formed. To determine whether the organ differences are associated with differences in the expression of organ identity genes, cDNA clones representing the putative homologs of B and C function MADS box genes were isolated and used in an in situ hybridization analysis. The expression of RAD1 and RAD2 (two different DEFICIENS homologs) in males and females was confined to the stamen whorl; the lack of expression in the second, inner perianth whorl correlated with the sepaloid nature of the inner whorl of perianth segments. Expression of RAP1 (a PLENA homolog) occurred in the carpel and stamen whorls in very young flower primordia from both males and females. However, as soon as the inappropriate set of organs ceased to develop, RAP1 expression became undetectable in those organs. The absence of expression of RAP1 may be the cause of the arrest in organ development or may be a consequence.     

SCIADOPITOPHYLLUM CANADENSE GEN. ET SP. NOV.: A NEW CONIFER FROM WESTERN ALBERTA

A new Sciadopitys-like conifer is described on the basis of compression fossils of shoots and leaves found at the Smokey Tower locality in western Alberta. The specimens consist of long, strap-like leaves attached in apparent whorls and subtended by groups of scale leaves. Other scale leaves are borne in loose spirals on the shoots between whorls. These specimens represent the first record of Sciadopitys-like foliage from western North America. Comparisons are made with extant and extinct species of the genus Sciadopitys (Siebold and Zuccarini, 1841) and with the widely distributed fossil genus Sciadopitytes (Goeppert and Menge, 1883).     

New Sphenophyllum plant from the Upper Devonian of Zhejiang Province,China

Sphenophyllum was an important and long-surviving sphenopsid genus in the Paleozoic floras, with a worldwide distribution. A new species, Sphenophyllum changxingense sp. nov., is described from the Upper Devonian Wutong Formation of Changxing County, Zhejiang Province, China. This plant is characterized by two orders of slender axes and wedge-shaped leaves borne in whorls. The axes bear short spines and show longitudinal ridges and furrows on surface. Three to eight isophyllous leaves, with one, two, or no second-order axes, are attached at each node of first-order axes. Leaves bear spines and show a bilobate morphology; the two leaf lobes divide distally to form several marginal segments, each segment with a leaf vein. Sphenophyllum changxingense represents an early and primitive species within the genus, in light of the absence of heterophylly and specialized hook-like leaves. Like some Carboniferous and Permian species, it appears to have formed dense mats with mutually supportive axes. This plant adds to the known diversity of early sphenopsids in the Late Devonian.     

A novel lycopsid from the Upper Devonian of Jiangsu, China

A new lycopsid, Monilistrobus yixingensis gen. et sp. nov., is described from the Wutung Formation (Famennian, Upper Devonian) of Jiangsu, China. The plant has many features typical of other Upper Devonian lycopsids, including dichotomous branching, helically arranged obovate expanded leaf bases, linear leaves with spiny appendages along the lateral margins, sporophylls widened proximally, and one elliptical sporangium attached to the adaxial surface. The most distinctive novel feature of the plant is that the modified sporophylls are arranged tightly into fertile zones or cone-like structures that are separated by lengths of axes with more lax sterile microphylls only: therefore the cone-like structures are strung along the branches like beads on a necklace.