Cycad Leaf Research Articles

Disparity of cycad leaves dispels the living fossil metaphor

The living fossil metaphor is tightly linked with the cycads. This group of gymnosperms is supposed to be characterised by long-term morphological stasis, particularly after their peak of diversity and disparity in the Jurassic. However, no formal test of this hypothesis exists. Here, we use a recent phylogenetic framework and an improved character matrix to reconstruct the Disparity Through Time for cycad leaves using a Principal Coordinate Analysis and employing Pre-Ordination Ancestral State Reconstruction to test the impact of sampling on the results. Our analysis shows that the cycad leaf morsphospace expanded up to the present, with numerous shifts in its general positioning, independently of sampling biases. Moreover, they also show that Zamiaceae expanded rapidly in the Early Cretaceous and continued to expand up to the present, while now-extinct clades experienced a slow contraction from their peak in the Triassic. We also show that rates of evolution were constantly high up to the Early Cretaceous, and then experienced a slight decrease in the Paleogene, followed by a Neogene acceleration. These results show a much more dynamic history for cycads, and suggest that the ‘living fossil’ metaphor is actually a hindrance to our understanding of their macroevolution.

Plant Megafossils, Palynomorphs, and Paleoenvironment from the Late Middle to Late Eocene Burnaby Mountain Flora, Huntingdon Formation, British Columbia, Canada

Premise of research. The Eocene fossil flora of the area around Vancouver, British Columbia, is poorly known despite work beginning in the 1890s to 1920s. The floristic character of the previously unstudied Burnaby Mountain flora from the Huntingdon Formation in British Columbia is reconstructed using plant megafossils and palynology. This site offers insight into the terrestrial vegetation and paleoclimate during the late middle to late Eocene of the Pacific Northwest of North America in a coastal setting during a global cooling trend.Methodology. Megaflora and microflora were identified, and the combined flora was compared with that of coeval floras from northwestern Washington. Paleoclimate was reconstructed from leaf morphology using the Climate Leaf Analysis Multivariate Program, leaf margin analysis, and leaf area analysis. A probabilistic nearest-living-relative approach was used to reconstruct paleoclimate independently of leaf morphology, using taxonomic identifications from both mega- and microfossils. These data were combined in an ensemble approach.Pivotal results. The Burnaby Mountain fossil flora is late middle Eocene to late Eocene in age and shares key plant taxa with the coeval Upper Ravenian flora of the Puget Group and the upper Chumstick Formation of northwestern Washington. The fossil flora contained a mix of subtropical and temperate forest elements, including rare palm and possible cycad leaf fragments, rare conifer pollen, and a diversity of broad-leaved trees.Conclusions. The reconstructed paleoclimate suggests humid warm-temperate to marginally subtropical conditions in coastal British Columbia during the late middle Eocene to late Eocene. An ensemble paleoclimate approach provided a most parsimonious mean annual temperature estimate of 16.2°C±3.1°C for the Burnaby Mountain fossils and mean annual precipitation of 134±56 cm. A modern climatic analog is present on the East Coast of the United States in North Carolina, where palms are part of the native flora.

Leaf nutrients of two Cycas L. species contrast among in situ and ex situ locations

An understanding of leaf nutrient relations is required for tree conservation and horticulture success. The study of cycad leaf nutrient dynamics has expanded in recent years, but direct comparisons among reports remains equivocal due to varying sampling protocols. We used Cycas micronesica K.D. Hill and Cycas nongnoochiae K.D. Hill trees to determine the influence on leaf nutrient concentrations of in situ versus ex situ locations and orientation of leaves within the tree canopy. Nitrogen, phosphorus, and potassium concentrations of leaves from ex situ plants exceeded those from in situ plants, and the differences were not explained by soil nutrient differences. Calcium concentrations of leaves varied among the site pairs, with differences primarily explained by soil calcium. Magnesium concentrations of leaves were not different among all location pairs even though soil magnesium concentrations varied among the sites more than any of the other elements. Differences in leaf macronutrient concentrations among four C. micronesica provenances were minimal when grown in a common garden. Lateral orientation of leaves did not influence any of the essential elements for either of the species. These findings indicate that the lateral orientation of cycad leaves does not influence leaf nutrient concentrations, leaf nutrient relations of cycad plants in managed ex situ settings do not align with leaf nutrient relations in habitat, and these differences are not explained by soil nutrition for most elements. We suggest that leaf nutrient concentrations should be determined in all niche habitats within the geographic range of a cycad species in order to fully understand the leaf physiology of each species.

Genomic characterization of cycad leaf necrosis virus, the first badnavirus identified in a gymnosperm.

A previously undescribed badnavirus was isolated from Zamia fischeri showing symptoms of chlorosis, necrosis, and ringspot. The virus has bacilliform virions 30nm in diameter and averaging 120nm in length. The viral genome is 9227bp in length and contains three open reading frames characteristic of members of the genus Badnavirus. The largest open reading frame (ORF3) encodes a putative polyprotein, with predicted domains including zinc finger, aspartic protease, reverse transcriptase (RT) and RNase H. The virus is tentatively named "cycad leaf necrosis virus" (CLNV). Within the genus Badnavirus, CLNV was most closely related to sugarcane bacilliform Guadeloupe D virus (FJ439817), sharing 69% identity at the nucleotide level in the RT + RNase H region. This virus is the first badnavirus reported to infect cycads, and it has the largest genome among the currently characterized badnaviruses.

Incident Light and Leaf Age Influence Leaflet Element Concentrations of Cycas micronesica Trees

The need for improved knowledge on conservation and management of cycad species has generated recent interest in compiling a database on leaf nutrient concentrations. However, the sampling protocols have not been consistent among reports and the influences of some plant and habitat traits on the plasticity of cycad leaf nutrient concentrations has not been adequately determined. We used Cycas micronesica K.D. Hill trees to determine the role of incident light level and leaf age on leaflet content of 11 essential elements. Shade leaves exhibited increased mass-based concentration for nitrogen, phosphorus, and potassium above that of sun leaves. Shade leaves exhibited decreased area-based concentration for all of the macro- and micronutrients below that of sun leaves. Mass-based concentration of nitrogen, phosphorus, and potassium decreased with leaf age, and that of calcium, magnesium, iron, manganese, and zinc increased with leaf age. These findings indicate the relative leaf age and the amount of shade or incident light at the leaf level must be recorded and reported for leaf tissue studies in cycads in order to reduce ambiguity and ensure repeatability.

Distribution of Elements along the Rachis of Cycas micronesica Leaves: A Cautionary Note for Sampling Design

Cycas micronesica K.D. Hill trees on the island of Yap were used to determine the influence of position along the leaf rachis on macro- and micro-nutrient concentrations and how leaf age affected the results. The outcomes revealed improvements to sampling protocols for future cycad leaf research. The concentration of every element except carbon and copper was influenced by leaflet position along the rachis. Most elements exhibited similar patterns for the oldest and youngest leaves on a tree, but the influence of position along the rachis for nitrogen, phosphorus, calcium, zinc, and boron was highly contrasting for old versus young leaves. The elements with the greatest variability along the rachis were potassium, phosphorus, manganese, and zinc, with the difference in basal and terminal leaflets as great as four-fold. Sampling leaflets at one position on a cycad leaf may generate inaccurate elemental concentration results for most essential nutrients other than carbon and copper. We have added position of sampled leaflets within leaves as a mandatory component of what is recorded and reported for future cycad leaf tissue analyses. Leaflets that span the full length of the rachis should be included in cycad leaf samples that are collected for tissue analysis.

Cracking the omega code: hydraulic architecture of the cycad leaf axis.

The leaf axis of members of the order Cycadales ('cycads') has long been recognized by its configuration of independent vascular bundles that, in transverse section, resemble the Greek letter omega (hence the 'omega pattern'). This provides a useful diagnostic character for the order, especially when applied to paleobotany. The function of this pattern has never been elucidated. Here we provide a three-dimensional analysis and explain the pattern in terms of the hydraulic architecture of the pinnately compound cycad leaf. The genus Cycas was used as a simple model, because each leaflet is supplied by a single vascular bundle. Sequential sectioning was conducted throughout the leaf axis and photographed with a digital camera. Photographs were registered and converted to a cinematic format, which provided an objective method of analysis. The omega pattern in the petiole can be sub-divided into three vascular components, an abaxial 'circle', a central 'column' and two adaxial 'wings', the last being the only direct source of vascular supply to the leaflets. Each leaflet is supplied by a vascular bundle that has divided or migrated directly from the closest wing bundle. There is neither multiplication nor anastomoses of vascular bundles in the other two components. Thus, as one proceeds from base to apex along the leaf axis, the number of vascular bundles in circle and column components is reduced distally by their uniform migration throughout all components. Consequently, the distal leaflets are irrigated by the more abaxial bundles, guaranteeing uniform water supply along the length of the axis. The omega pattern exemplifies one of the many solutions plants have achieved in supplying distal appendages of an axis with a uniform water supply. Our method presents a model that can be applied to other genera of cycads with more complex vascular organization.

Praeornis sharovi Rautian, 1978 a fossil feather from the early Late Jurassic of Kazakhstan

Since its original description as a feather belonging to a basal bird, the phylogenetic position of Praeornis sharovi was debated. It was considered as belonging to a bird, a cycad leaf, or as a ‘transitional’ integumentary structure between reptile scales and bird feathers. Recently, a basal enantiornithine bird was collected in Early Cretaceous beds of Brazil. This specimen shows very well-preserved rachis-dominated tail feathers with a very thick rachis and thick and rigid barbules. These features are present in Praeornis, suggesting that this fossil may be interpreted as the tail feather of a basal bird. In this way, Praeornis constitutes one of the oldest records of rachis-dominated feathers in the world. Rachis-dominated tail feathers, including that of Praeornis appear to be rigid paired structures not performed for aerodynamical purposes, suggesting that may be important in body balance.

Molecular characterization of two badnavirus genomes associated with Canna yellow mottle disease

Members of the genus Badnavirus have a single non-covalently closed circular double-stranded DNA genome of 7.2–9.2kb. The genome encodes three open reading frames (ORFs) on the positive DNA strand. Canna yellow mottle virus (CaYMV) is a badnavirus that has been described as the etiological cause of yellow mottle disease in canna, although only a 565bp fragment of the genome has been previously reported from cannas. In this report, concentrated virions were recovered from infected canna plants and nucleic acids were extracted. Two full-length sequences represent two badnavirus genomes were recovered and were determined to be 6966bp and 7385bp in length. These DNAs represent a virus strain belonging to Canna yellow mottle virus and a novel species tentatively termed Canna yellow mottle associated virus. Phylogenetic analysis indicates that these two viruses are closely related to sugarcane bacilliform GD virus, pineapple bacilliform comosus virus, banana streak MY virus, and cycad leaf necrosis virus. We also showed naturally grown canna plants to be frequently co-infected by these two badnaviruses along with a potyvirus, Canna yellow streak virus.

Patchy distribution and varied habitats of Macrozamia lucida cycads explained by constancy in a key environmental variable

Identifying the influence of environmental variables on the spatio-temporal distribution of organisms remains a central goal of ecology. Although environmental limits to the geographic range of species have been investigated, less information is available regarding the influence of environmental variables limiting species to discrete patches within their geographic range. Identifying environmental variables of importance to the species in question is further complicated where the patches appear to be distributed across a variety of outwardly dissimilar habitats. We assessed the influence of environmental variables on the patchy distribution of Macrozamia lucida L.A.S. Johnson (Zamiaceae) cycads endemic to south-east Queensland, Australia. Here we show that patches of M. lucida are consistently associated with elevated moisture, despite the patches being located in contrasting habitats and topographic positions. Our survey results across five field sites demonstrate that both cycad abundance and adult cycad condition (estimated from measures of adult cycad leaf production and reproductive output) are positively correlated with elevated moisture, as measured by mean minimum relative humidity at the plant–soil interface. The historical biogeography of the genus Macrozamia may be intimately linked with the contraction and fragmentation of Australia’s mesic biomes during the Miocene. It is plausible that current distribution pattern of M. lucida, restricted to discrete patches that meet the species’ environmental requirements, reflects a contraction and fragmentation within its range from a more regular historical distribution under more favourable climatic regimes.

Does cycad aulacaspis scale (Aulacaspis yasumatsui, Hemiptera: Diaspididae) play a direct role in causing soil phytotoxicity?

Cycad aulacaspis scale (CAS, Aulacaspis yasumatsui, Hemiptera: Diaspididae) was accidentally introduced to Guam in 2003, and has caused acute mortality of the dominant, endemic forest tree Cycas micronesica. A phytotoxic legacy in the soils beneath cycad trees killed by CAS over a period of about three years has been demonstrated. The origin of the toxicity may be large quantities of CAS-encrusted cycad leaf litter. We explore the possibility that a major contribution to this toxic legacy may come from the scale insects, not just from the plant material.

First beetle elytra, abdomen (Coleoptera) and a mine trace from Lunz (Carnian, Late Triassic, Lunz‐am‐See, Austria) and their taphonomical and evolutionary aspects

Abstract: Four new species of Permosynidae are described: Platycrossus caroli Ponomarenko in Meller et al. sp. nov., Hydrobiites handlirschi Ponomarenko in Meller et al. sp. nov., Ademosyne polyzetete Ponomarenko in Meller et al. sp. nov. and Diarcuipenna bennettitophila Ponomarenko in Meller et al. sp. nov. Together with one polyphagous abdomen, they represent the first beetle remains from the Lunz formation, known for its richness in bennettite and cycad leaf remains and also for the bennettite reproductive organs. Furthermore, the first insect mine trace on a Nilssonia leaf segment is described, which is one of the most ancient linear mine traces. The rareness of insect remains in the plant‐bearing layers of the Lunz formation is still an enigma from the taphonomical–chemical point of view. The palaeoentomological and palaeobotanical considerations tentatively indicate a relationship between the rare occurrence of coniferous plants and the evolution or radiation of phytophagous insects during this time.

An Early Cretaceous zamiaceous cycad of South West Gondwana: Restrepophyllum nov. gen. from Patagonia, Argentina

The record of Cycadales in Patagonia begins in the Triassic and extends up to the Oligocene. In this region the group is highly diversified and includes several taxa represented by trunks, leaves and pollen cones. A new cycadalean genus and species, Restrepophyllum chiguoides, form the Aptian Anfiteatro de Ticó Formation, Santa Cruz province, Argentina, is described here. The fossil is a leaf compression with well-preserved cuticle. Its morphology, anatomy and ultrastructure are studied by means of light and electron microscopy. The leaf is lanceolate, serrate, and possesses a prominent midvein and decurrent laterals showing an open, simple or dichotomous venation. The leaf is hypostomatic, and the abaxial cuticle is thinner than the adaxial one. The stomata are irregularly distributed and indistinctly oriented between veins. They are monocyclic to imperfectly dicyclic (haplocheilic); the suprastomatal aperture is raised over the epidermis and the guard cells are sunken. Scattered trichomes and crystalliferous idioblasts are also present. The cuticle is composed of three layers: the outer and inner layers are lamellate, while the middle one is granulate. This new cycad leaf is compared with similar fossil leaves from Gondwana and Europe/North America, and also with similar extant cycad leaves. Based on the general morphology and the main characters of the cuticle, R. chiguoides is assigned to the family Zamiaceae; moreover it is more closely related to the living Zamia ( Chigua) restrepoi (D. Stevenson) Lindstrom than to any other member of the Cycadales. Paleophytogeographic evidence suggests a South American origin of Zamia/Chigua and a further migration to northern latitudes. This new type of leaf also suggests the putative existence of a Chigua clade that may be traced back to the Early Cretaceous when two cycadalean families, Zamiaceae and Stangeriaceae, were already well-established in Patagonia.

Cycads from the Triassic of Antarctica: Permineralized Cycad Leaves

Permineralized cycad petioles and/or rachides with associated pinnae are described from two Triassic localities in the Queen Alexandria Range, central Transantarctic Mountains, Antarctica. Petiole‐rachides display an inverted‐omega‐shaped arrangement of vascular bundles typical of most genera of extant Cycadales and exarch primary xylem that link them to the modern order. Pinnae associated with the Antarctic petiole‐rachides are thin, with regularly spaced vascular bundles. They are similar to those of extant Zamia and most other genera of extant Cycadales, whose pinnae lack midribs. Other Mesozoic fossil cycads (e.g., Charmorgia, Lyssoxylon, Lioxylon) have endarch petiole vascular bundles that in some cases were previously considered more similar to those of Bennettitales than those of Cycadales. We suggest, however, that the endarch xylem of these taxa is typical of Cycadales because in extant cycads, the protoxylem changes from endarch to exarch within the base of the petiole. Evolution of cycad leaf form is reviewed based on evidence from the fossil record.

Cloning and Expression of a Mannose-Binding Jacalin-Related Lectin from Leaves of Japanese Cycad (Cycas revolutaThunb.)

Cycad leaf lectin (CRLL), a mannose-recognizing jacalin-related lectin (mJRL), was first cloned as a gymnosperm lectin and expressed. The cDNA sequence of CRLL (DDBJ, accession no. AB198328), coding 291 amino acid residues, has a tandem repeat of about 150 amino acids divided into N- and C-terminal domains as Japanese chestnut mJRL. Sequence alignment showed deletion and insertion of the sequence, and its putative carbohydrate-binding sites showed some differences from other JRLs. PCR analysis showed that this lectin was expressed in the cycad leaf but not in the root or seed. Recombinant CRLL (rCRLL) was expressed in Escherichia coli and purified by affinity chromatography after refolding procedures. Properties of active rCRLL appeared to be almost the same as those of native CRLL.

Cuticular characters adapted to volcanic stress in a new Cretaceous cycad leaf from Patagonia, Argentina. Considerations on the stratigraphy and depositional history of the Baqueró Formation

The cuticle of a new cycad, Pseudoctenis ornata Archangelsky et al., sp. nov. is described and discussed in relation to the physical paleoenvironment in which the plant lived. The specimens occur in the Early Cretaceous Baqueró Formation, near Estancia El Verano in the Santa Cruz Province, Argentina. A detailed stratigraphic section records four facies, namely (1) fluvial channel, (2) flood plain, (3) lacustrine, and (4) flat and extended plains. A detail of each facies is provided. Pseudoctenis cuticles are found in the flood plain facies; the other components of the plant association are Gleichenites, Araucaria and Taeniopteris. The depositional history of this succession is related to a braided river that periodically received volcanic ash. Plants grew until complete burial by ash. Leaves of Pseudoctenis are pinnate, hypostomatic, with polycyclic stomata that form ill-defined rows. Abundant papillae and hair bases are present, especially on the lower cuticle. Comparisons are made with other Pseudoctenis species found in the same formation, and in other regions of the world. It is suggested that the paleoenvironment had a strong influence on the vegetation, especially the ash fall, and that it may have played a role in the formation of xeromorphic structures that characterize several gymnosperms present in this stratigraphic unit, including Pseudoctenis ornata. The lack of burning features on the cuticle suggests a cold ash fall. This fall probably was responsible for changes in edaphic patterns and atmospheric conditions that governed the growth and distribution of plant communities during the deposition of the Baqueró Formation.

A New Pinnate Cycad Leaf from the Upper Triassic Chinle Formation of Arizona

A recently discovered Upper Triassic leaf shows many of the characters of the leaves of the living cycads and is assigned to the Cycadales. The leaf, however, does not match any previously described fossil or living cycad leaf in all features and is assigned to a new genus and species, Aricycas paulae. The leaf bears oppositely arranged linear-lanceolate pinnae with pointed apices and contracted bases Pinnae contain prominent midribs and anastomosing lateral veins which end freely near the margins. The ordinary epidermal cells are rectangular with straight to slightly curving side walls and haplocheilic stomata. Pinnae in this leaf are apparently wrinkled and shriveled and in some cases fell before the parent leaves were shed, as is typical of the living cycads. It appears that this character has never been reported in fossil cycad leaves. Since A. paulae is Late Carnian age, it is one of the oldest known cycad leaves and thus supports the suggestion by Delevoryas (1982) that there is as much evidence that plants with compound leaves could have given rise to the Cycadales as plants with entire leaves have.