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Concept: Cycas


The Asian cycads are mostly allopatric, distributed in small population sizes. Hybridization between allopatric species provides clues in determining the mechanism of species divergence. Horticultural introduction provides the chance of interspecific gene flow between allopatric species. Two allopatrically eastern Asian Cycas sect. Asiorientales species, C. revoluta and C. taitungensis, which are widely distributed in Ryukyus and Fujian Province and endemic to Taiwan, respectively, were planted in eastern Taiwan for horticultural reason. Higher degrees of genetic admixture in cultivated samples than wild populations in both cycad species were detected based on multilocus scans by neutral AFLP markers. Furthermore, bidirectional but asymmetric introgression by horticultural introduction of C. revoluta is evidenced by the reanalyses of species associated loci, which are assumed to be diverged after species divergence. Partial loci introgressed from native cycad to the invaders were also detected at the loci of strong species association. Consistent results tested by all neutral loci, and the species-associated loci, specify the recent introgression from the paradox of sharing of ancestral polymorphisms. Phenomenon of introgression of cultivated cycads implies niche conservation among two geographic-isolated cycads, even though the habitats of the extant wild populations of two species are distinct.

Concepts: Biology, Taiwan, Gene flow, Cycad, Cycas, Cycas revoluta


Crypticerya zeteki (Cockerell, 1914) (Hemiptera: Coccomorpha: Monophlebidae) is recorded for the first time from Brazil and Maconellicoccus hirsutus (Green, 1908) (Hemiptera: Coccomorpha: Pseudococcidae) is recorded for the first time from the state of Maranhão, Brazil. Both species were collected from branches, leaves and fruits of various fruit trees in the municipalities of São José de Ribamar, São Luís and Paço do Lumiar, Maranhão, Brazil. Crypticerya zeteki was collected on Citrus spp. (Rutaceae), Cocos nucifera (L.) (Arecaceae), Cycas revoluta L. (Cycadaceae), Malpighia punicifolia L. (Malpighiaceae), Mangifera indica L. (Anacardiaceae), Musa paradisiaca L. (Musaceae) and Theobroma grandiflorum Schum (Malvaceae), all first records for this species. Maconellicoccus hirsutus was collected on Spondias tuberosa Arruda (Anacardiaceae) and M. punicifolia L. (Malpighiaceae), both new records for this species.

Concepts: Arecaceae, Fruit, Seed, Mangifera, Banana, Cycad, Cycas, Cycas revoluta


Most knowledge concerning Mesozoic Era floras has come from compression fossils. This has been augmented in the last 20 years by rarer permineralized material showing cellular preservation. Here, we describe a new genus of anatomically preserved gymnosperm seed from the Callovian-Oxfordian (Jurassic) Oxford Clay Formation (UK), using a combination of traditional sectioning and synchrotron radiation X-ray micro-tomography (SRXMT). Oxfordiana motturii gen. et sp. nov. is large and bilaterally symmetrical. It has prominent external ribs, and has a three-layered integument comprising: a narrow outer layer of thick walled cells; a thick middle parenchymatous layer; and innermost a thin fleshy layer. The integument has a longitudinal interior groove and micropyle, enveloping a nucellus with a small pollen chamber. The large size, bilateral symmetry and integumentary groove demonstrate an affinity for the new species within the cycads. Moreover, the internal groove in extant taxa is an autapomorphy of the genus Cycas, where it facilitates seed germination. Based upon the unique seed germination mechanism shared with living species of the Cycadaceae, we conclude that O. motturii is a member of the stem-group lineage leading to Cycas after the Jurassic divergence of the Cycadaceae from other extant cycads. SRXMT-for the first time successfully applied to fossils already prepared as slides-reveals the distribution of different mineral phases within the fossil, and allows us to evaluate the taphonomy of Oxfordiana. An early pyrite phase replicates the external surfaces of individual cells, a later carbonate component infilling void spaces. The resulting taphonomic model suggests that the relatively small size of the fossils was key to their exceptional preservation, concentrating sulfate-reducing bacteria in a locally closed microenvironment and thus facilitating soft-tissue permineralization.

Concepts: Seed, Germination, Fossil, Ovule, Gymnosperm, Mesozoic, Cycad, Cycas


Chilades pandava (Lepidoptera: Lycaenidae) larval food quality was studied to determine its influence on adult life history traits. A wild population from Cycas nongnoochiae (Cycadales: Cycadaceae) endemic habitat behaved similarly to the population collected from a garden setting. Cycas micronesica, Cycas revoluta, and Cycas seemannii leaves were used as high-quality food, whereas C nongnoochiae, Cycas taitungensis, and Cycas condaoensis leaves were used as low-quality food. The daily oviposition rate was not influenced by food quality, but longevity and lifetime fecundity of females were increased by high-quality larval food. These results indicate that in situ Cycas species impose a physiological constraint on the genetic capacity to produce offspring by C pandava. The removal of that constraint by high-quality novel Cycas species may be one reason this butterfly can increase in population rapidly after an invasion event and express greater herbivory of Cycas species within invaded regions.

Concepts: Biology, Lepidoptera, Butterfly, Cycad, Cycas, Cycas revoluta, Cycas circinalis, Cycas taitungensis


Dramatic crustal deformation and river incision in Southwest China induced by the Indo-Asian collision have long been argued to contribute to the complicated landscapes, heterogeneous environment and abundant biodiversity in this region. However, biological impacts in promoting intraspecific phylogeographical subdivision and divergence along the Red River Fault zone (RRF) remain poorly understood. To investigate the possible biological effects of tectonic movements and environment variations within the RRF, the phylogeography of Cycas dolichophylla-an endemic but widely distributed Cycas in Southwest China and North Vietnam along the RRF were carried out based on four chloroplast DNA intergenic spacers (cpDNA), three nuclear DNA sequences (nDNA) and 16 simple sequence repeat variations (SSR). Two different phylogeographical patterns were detected: a Southwest-Northeast break across the RRF disclosed by chlorotypes and a China-Vietnam separation revealed by SSR. A Bayesian skyline plot from cpDNA data demonstrated a historical increasing, but a recent declining, dynamic in population size during the Pleistocene. Consequently, we infer it is the local environmental variation during Cenozoic that contributed to the complex landscape and microclimate mosaics, facilitating speciation and divergence of C. dolichophylla. Subsequently, the Quaternary climatic fluctuations coupled with human activities profoundly influenced the genetic structure and demographic history of this species.

Concepts: DNA, Genetics, Evolution, Eukaryote, Species, Chloroplast, Yunnan, Cycas


In angiosperms, gene conversion has been known to reduce the mutational load of plastid genomes (the plastomes). Particularly, more frequent gene conversions in inverted repeat (IR) than in single copy (SC) regions result in contrasting substitution rates between these two regions. However, little has been known about the effect of gene conversion in the evolution of gymnosperm plastomes. Cycads (Cycadophyta) are the second largest gymnosperm group. Evolutionary study of their plastomes is limited to the basal cycad genus, Cycas. In this study, we addressed three questions. (1) Do the plastomes of other cycad genera evolve slowly as previously observed in the plastome of Cycas taitungensis? (2) Do substitution rates differ between their SC and IR regions? And (3) Does gene conversion occur in the cycad plastomes? If yes, is it AT-biased or GC-biased? Plastomes of eight species from other eight genera of cycads were sequenced. These plastomes are highly conserved in genome organization. Excluding ginkgo, cycad plastomes have significantly lower synonymous and non-synonymous substitution rates than other gymnosperms, reflecting their evolutionary stasis in nucleotide mutations. In the IRs of cycad plastomes, the reduced substitution rates and GC-biased mutations are associated with a GC-biased gene conversion (gBGC) mechanism. Further investigations suggest that in cycads, gBGC is able to rectify plastome-wide mutations. Therefore, this study is the first to uncover the plastomic gBGC in seed plants. We also propose a gBGC model to interpret the dissimilar evolutionary patterns as well as the compositionally biased mutations in the SC and IR regions of cycad plastomes.

Concepts: DNA, Gene, Genetics, Evolution, Plant, Gymnosperm, Cycad, Cycas


Transglycosylation (TG) activity of a family GH18 chitinase from the cycad, Cycas revoluta, (CrChiA) was modulated by removing or introducing a tryptophan side chain. The removal from subsite +3 through mutation of Trp168 to alanine suppressed TG activity, while introduction into subsite +1 through mutation of Gly77 to tryptophan (CrChiA-G77W) enhanced TG activity. The crystal structures of an inactive double mutant of CrChiA (CrChiA-G77W/E119Q) with one or two N-acetylglucosamine residues occupying subsites +1 or +1/+2, respectively, revealed that the Trp77 side chain was oriented toward +1 GlcNAc to be stacked with it face-to-face, but rotated away from subsite +1 in the absence of GlcNAc at the subsite. Aromatic residues in the aglycon-binding site are key determinants of TG activity of GH18 chitinases.

Concepts: DNA, Modulation, Module, Website, Cycad, N-Acetylglucosamine, Cycas, Cycas revoluta


A class V (GH family 18) chitinase from the cycad, Cycas revoluta (CrChiA) is a plant chitinase, that was reported to possess efficient transglycosylation (TG) activity. We solved the crystal structure of CrChiA and compared it with those of class V chitinases from Nicotiana tabaccum (NtChiV) and Arabidopsis thaliana (AtChiC), which do not efficiently catalyze the TG reaction. All three chitinases had a similar (α/β)8 barrel fold with an (α+β) insertion domain. In the acceptor-binding site (+1, +2, and +3) of CrChiA, the Trp168 side chain was found to stack face-to-face with the +3 sugar. However, this interaction was not found in the identical regions of NtChiV and AtChiC. In the DxDxE motif, which is essential for catalysis, the carboxyl group of the middle Asp (Asp117) was always oriented toward the catalytic acid Glu119 in CrChiA, whereas the corresponding Asp in NtChiV and AtChiC was oriented toward the first Asp. These structural features of CrChiA appeared to be responsible for the efficient TG activity. When inhibitor (allosamidin) binding ability was evaluated using isothermal titration calorimetry, binding free energy changes of the three chitinases were found to be similar to each other; -9.5 ~ -9.8 kcal/mol. However, solvation and conformational entropy changes in CrChiA were marledly different from those in NtChiV and AtChiC, but similar to those in chitinase A from Serratia marcescens (SmChiA), which also exhibits significant TG activity. These results have provided insights into the molecular mechanism underlying the TG reaction and the molecular evolution from bacterial chitinases to plant class V chitinases. This article is protected by copyright. All rights reserved.

Concepts: Catalysis, Arabidopsis thaliana, Arabidopsis, Solid, Entropy, Copyright, Cycad, Cycas


Cycas is the most widespread and diverse genus among the ancient cycads, but the extant species could be the product of late Miocene rapid radiations. Taxonomic treatments to date for this genus are quite controversial, which makes it difficult to elucidate its evolutionary history. We cloned 161 genomic ITS sequences from 31 species representing all sections of Cycas. The divergent ITS paralogs were examined within each species and identified as putative pseudogenes, recombinants and functional paralogs. Functional paralogs were used to reconstruct phylogenetic relationships with pseudogene sequences as molecular outgroups, since an unambiguous ITS sequence alignment with their closest relatives, the Zamiaceae, is unachievable. A fully resolved and highly supported tree topology was obtained at the section level, with two major clades including six minor clades. The results fully supported the classification scheme proposed by Hill (2004) at the section level, with the minor clades representing his six sections. The two major clades could be recognised as two subgenera. The obtained pattern of phylogenetic relationships, combined with the different seed dispersal capabilities and paleogeography, allowed us to propose a late Miocene rapid radiation of Cycas that might have been promoted by vicariant events associated with the complex topography and orogeny of South China and adjacent regions. In contrast, transoceanic dispersals might have played an important role in the rapid diversification of sect. Cycas, whose members have evolved a spongy layer in their seeds aiding water dispersals.

Concepts: DNA, Gene, Evolution, Biology, Species, Phylogenetics, Cycad, Cycas


Several individuals of the Caribbean Zamia clade and other cycad genera were used to identify single copy nuclear genes for phylogeographic and phylogenetic studies in Cycadales. Two strategies were employed to select target loci: 1) a tblastX search of Arabidopsis conserved ortholog sequence (COS) set and, 2) a tblastX search of Arabidopsis-Populus-Vitis-Oryza Shared Single Copy genes (APVO SSC) against the EST Zamia databases in Genbank. From the first strategy, 30 loci were selected, and from the second, 16 loci. In both cases the matching Genbank accessions of Zamia were used as a query for retrieving highly similar sequences from Cycas, Picea, Pinus species or Ginko biloba. After retrieving and aligning all the sequences in each locus, intron predictions were completed to assist in primer design. PCR was carried out in three rounds to detect paralogous loci. A total of 29 loci were successfully amplified as a single band of which 20 were likely single copy loci. These loci showed different diversity and divergence levels. A preliminary screening allowed us to select 8 promising loci (40S, ATG2, BG, GroES, GTP, LiSH, PEX4 and TR) for the Z. pumila complex and 4 loci (COS26, GroES, GTP and HTS) for all other cycad genera. This article is protected by copyright. All rights reserved.

Concepts: Polymerase chain reaction, Species, Ginkgo biloba, Copyright, Ginkgo, Gymnosperm, Cycad, Cycas