Zirabia cylindrica comb. nov. provides evidence of Doyleales in the Jurassic.

PREMISE: In recent years, Doyleales have played an important role in the discussion and exploration of seed plant relationships and the origin of angiosperms. This order comprises a diversity of recently described genera with cupule-bearing compound ovulate cones recovered from Early Cretaceous deposits in North America and Asia. Their relatively late appearance in the fossil record, stratigraphically near the appearance of angiosperms in the Early Cretaceous, has been noteworthy. Here, we report a new genus of Doyleales, Zirabia gen. nov. from the Early Jurassic of Iran, that was originally described as the ginkgophyte Karkenia. METHODS: We reinvestigated material previously assigned to Karkenia cylindrica from the Lower Jurassic of the Zirab locality, Alborz Mountains, northern Iran. RESULTS: The studied specimen features a main axis bearing helically to irregularly arranged bract-cupule complexes, each composed by a long laminar bract subtending and sheathing a cupule stalk that bears a single-seeded cupule with a dorsal protrusion. The morphological features of this taxon do not conform with those of Karkenia, and suggest affinities with Doyleales rather than Ginkgoales. Within Doyleales, this fossil has a unique combination of characters indicating that it is a new genus within the order; thus, a new combination is erected, Zirabia cylindrica. CONCLUSIONS: Our results indicate that Doyleales is significantly older than previously thought, with their stratigraphic range now extending from the Lower Jurassic to the Cretaceous. The Early Jurassic occurrence of Doyleales provides important data on the emergence of seed-enclosing structures seen in seed plants throughout the Mesozoic.

Cretaceous pollen cone with three-dimensional preservation sheds light on the morphological evolution of cycads in deep time.

The Cycadales are an ancient and charismatic group of seed plants. However, their morphological evolution in deep time is poorly understood. While molecular divergence time analyses estimate a Cretaceous origin for most major living cycad clades, much of the extant diversity is inferred to be a result of Neogene diversifications. This leads to long branches throughout the cycadalean phylogeny that, with few exceptions, have yet to be rectified by unequivocal fossil cycads. We report a permineralized pollen cone from the Campanian Holz Shale located in Silverado Canyon, CA, USA (c. 80 million yr ago). This fossil was studied via serial sectioning, SEM, 3D reconstruction and phylogenetic analyses. Microsporophyll and pollen morphology indicate this cone is assignable to Skyttegaardia, a recently described genus based on disarticulated lignitized microsporophylls from the Early Cretaceous of Denmark. Data from this new species, including a simple cone architecture, anatomical details and vasculature organization, indicate cycadalean affinities for Skyttegaardia. Phylogenetic analyses support this assignment and recover Skyttegaardia as crown-group Cycadales, nested within Zamiaceae. Our findings support a Cretaceous diversification for crown-group Zamiaceae, which included the evolution of morphological divergent extinct taxa with unique traits that have yet to be widely identified in the fossil record.

Origin of Equisetum: Evolution of horsetails (Equisetales) within the major euphyllophyte clade Sphenopsida.

PREMISE OF THE STUDY: Equisetum is the sole living representative of Sphenopsida, a clade with impressive species richness, a long fossil history dating back to the Devonian, and obscure relationships with other living pteridophytes. Based on molecular data, the crown group age of Equisetum is mid-Paleogene, although fossils with possible crown synapomorphies appear in the Triassic. The most widely circulated hypothesis states that the lineage of Equisetum derives from calamitaceans, but no comprehensive phylogenetic studies support the claim. Using a combined approach, we provide a comprehensive phylogenetic analysis of Equisetales, with special emphasis on the origin of genus Equisetum. METHODS: We performed parsimony phylogenetic analyses to address relationships of 43 equisetalean species (15 extant, 28 extinct) using a combination of morphological and molecular characters. KEY RESULTS: We recovered Equisetaceae + Neocalamites as sister to Calamitaceae + a clade of Angaran and Gondwanan horsetails, with the four groups forming a clade that is sister to Archaeocalamitaceae. The estimated age for the Equisetum crown group is mid-Mesozoic. CONCLUSIONS: Modern horsetails are not nested within calamitaceans; instead, both groups have explored independent evolutionary trajectories since the Carboniferous. Diverse fossil taxon sampling helps to shed light on the position and relationships of equisetalean lineages, of which only a tiny remnant is present within the extant flora. Understanding these relationships and early character configurations of ancient plant clades as Equisetales provide useful tests of hypotheses about overall phylogenetic relationships of euphyllophytes and foundations for future tests of molecular dates with paleontological data.

Developmental programmes in the evolution of Equisetum reproductive morphology: a hierarchical modularity hypothesis.

Background: The origin of the Equisetum strobilus has long been debated and the fossil record has played an important role in these discussions. The paradigm underlying these debates has been the perspective of the shoot as node-internode alternation, with sporangiophores attached at nodes. However, fossils historically excluded from these discussions (e.g. Cruciaetheca and Peltotheca ) exhibit reproductive morphologies that suggest attachment of sporangiophores along internodes, challenging traditional views. This has rekindled discussions around the evolution of the Equisetum strobilus, but lack of mechanistic explanations has led discussions to a stalemate. Scope: A shift of focus from the node-internode view to a perspective emphasizing the phytomer as a modular unit of the shoot, frees the debate of homology constraints on the nature of the sporangiophore and inspires a mechanism-based hypothesis for the evolution of the strobilus. The hypothesis, drawing on data from developmental anatomy, regulatory mechanisms and the fossil record, rests on two tenets: (1) the equisetalean shoot grows by combined activity of the apical meristem, laying down the phytomer pattern, and intercalary meristems responsible for internode elongation; and (2) activation of reproductive growth programmes in the intercalary meristem produces sporangiophore whorls along internodes. Conclusions: Hierarchical expression of regulatory modules responsible for (1) transition to reproductive growth; (2) determinacy of apical growth; and (3) node-internode differentiation within phytomers, can explain reproductive morphologies illustrated by Cruciaetheca (module 1 only), Peltotheca (modules 1 and 2) and Equisetum (all three modules). This model has implications - testable by studies of the fossil record, phylogeny and development - for directionality in the evolution of reproductive morphology ( Cruciaetheca - Peltotheca - Equisetum ) and for the homology of the Equisetum stobilus. Furthermore, this model implies that sporangiophore development is independent of node-internode identity, suggesting that the sporangiophore represents the expression of an ancestral euphyllophyte developmental module that pre-dates the evolution of leaves.