BackgroundThe extant Gnetales include three monotypic families, namely, Ephedraceae (Ephedra), Gnetaceae (Gnetum), and Welwitschiaceae (Welwitschia), all of which possess compound female cones that comprise a main axis and 1 to multiple pairs/whorls of bracts subtending a female reproductive unit or having lower pairs/whorls of bracts sterile. However, the evolutionary origin of such a reproductive architecture in Gnetales is controversial in the light of the competing anthophyte versus gnetifer hypotheses of seed plant relationships. Hence, macrofossils demonstrating the structure of compound female cones of the Gnetales should be important to decipher the early evolution of the order.ResultsA new ephedroid plant Chengia laxispicata gen. et sp. nov. is described from the Early Cretaceous Yixian Formation of western Liaoning, Northeast China. The fossil represents a part of a leafy shooting system with reproductive organs attached. The main shoot bears internodes and swollen nodes, from which lateral branches arise oppositely. Reproductive organs consist of female spikes terminal to twigs or axillary to linear leaves. Spikes are loosely arranged, having prominent nodes and internodes. Bracts of the spikes are decussately opposite and comprise 4—8 pairs of bracts. Each bract subtends an ellipsoid seed. Seeds are sessile, with a thin outer envelope and a distal micropylar tube.ConclusionsChengia laxispicata gen. et sp. nov. provides a missing link between archetypal fertile organs in the crown lineage of the Gnetales and compound female cones of the extant Ephedraceae. Combined with a wealth of Ephedra and ephedroid macrofossils from the Early Cretaceous, we propose a reduction and sterilization hypothesis that the female cone of the extant Ephedraceae may have stemmed from archetypal fertile organs in the crown lineage of the Gnetales. These have undergone sequentially intermediate links similar to female cones of Cretaceous Siphonospermum, Chengia, and Liaoxia by reduction and sterilization of the lower fertile bracts, shortenings of internodes and peduncles as well as loss of reproductive units in all inferior bracts. The basal family Ephedraceae including Ephedra of the extant Gnetales was demonstrated to have considerable diversity by the Early Cretaceous, so an emended familial diagnosis is given here. The Jehol Biota in Northeast China and adjacent areas contains a plethora of well-preserved macrofossils of Ephedra and ephedroids that show different evolutionary stages including primitive and derived characters of Ephedraceae, so Northeast China and adjacent areas may represent either the centre of origination or one of the centres for early diversification of the family.

In the present paper, an ephedroid macrofossil species from the Early Cretaceous Yixian Formation of western Liaoning of China is described as new to science: Ephedra multinervia Yang et Lin, sp. nov. This species has typical ephedroid morphology, e.g. the dichasial branching shoot system, swollen nodes, internodes having many fine longitudinal striations and opposite phyllotaxy. Ephedra multinervia has strap-shaped leaves with multiple dichotomizing veins and reduced female cones with a single pair of fertile bracts forming a cupule enclosing two inner seeds. Ephedra multinervia is similar to Ephedra archaeorhytidosperma Yang et al. and Ephedra hongtaoi Wang et Zheng in its reduced bi-ovulate female cone, but differs from the latter two species by the lengthy strap-like leaves bearing multiple parallel veins and its sessile female cones. A new evolutionary hypothesis of the gnetophytes is proposed based on a synthesis of reproductive morphology of macrofossils from the Early Cretaceous and modern representatives. A Chengia-like precursor might have given rise to the Gnetum-Welwitschia clade by diversification of leaf morphology and female reproductive organs. According to this new explanation, the Welwitschialike strap-like leaves with multiple parallel veins in E. multinervia result from convergence.

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BackgroundMale cones of modern Ephedraceae are compound and compact. No fossil evidence has so far been found to support an origin of the compact compound male cone from a hypothetical loosely-arranged shoot system.ResultsHere we describe a new macrofossil taxon, Eamesia chinensis Yang, Lin, Ferguson et Wang, gen. et sp. nov., from the Early Cretaceous of western Liaoning, northeastern China. It was an ephedroid shrub bearing male spikes terminal to twigs, but differs from modern Ephedraceae by its loosely-arranged male cones, the axillary male shoot consisting of an elongated synangiophore on which leaf-like foliar organs were inserted, and four sessile synangia terminal to the apex.ConclusionsThe morphology of this fossil suggests that the modern compact male cone of Ephedra was indeed derived from a once loosely-arranged shoot system, and the male reproductive unit originated from a once elongated axillary male shoot. This new fossil species thus provides a transitional link from the hypothetical ancestral shoot system to the modern compact morphology. Changes of habitat from closed humid forests to open dry deserts and shifts of the pollination syndrome may have acted as the driving forces behind this morphological evolution.

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Macrofossil evidence has demonstrated a first radiation of gnetophytes in the Early Cretaceous. However, the origin of the diversity of gnetophytes remains ambiguous because gnetalean macrofossils have rarely been reported from pre-Cretaceous strata. Here, we report a new putative gnetalean macrofossil reproductive shoot which possesses opposite phyllotaxy, long linear leaves more or less decurrent and having a prominent midvein and pedicled ovoid-ellipsoid and longitudinally striated chlamydosperms. Our new fossil is different from other known gnetalean macrofossils in the linear-lanceolate leaves with a midvein and pedicled chlamydosperms. As a result, we describe this new macrofossil reproductive shoot as new to science, i.e., Daohugoucladus sinensis gen. et sp. nov. Our new macrofossil displays additional morphological characters distinct from other known Mesozoic and modern gnetalean species and provides additional evidence of the origin and early evolution of female reproductive organs of gnetophytes.