From the Cover: Montsechia,an ancient aquatic angiosperm

The early diversification of angiosperms in diverse ecological niches is poorly understood. Some have proposed an origin in a darkened forest habitat and others an open aquatic or near aquatic habitat. The research presented here centers on Montsechia vidalii, first recovered from lithographic limestone deposits in the Pyrenees of Spain more than 100 y ago. This fossil material has been poorly understood and misinterpreted in the past. Now, based upon the study of more than 1,000 carefully prepared specimens, a detailed analysis of Montsechia is presented. The morphology and anatomy of the plant, including aspects of its reproduction, suggest that Montsechia is sister to Ceratophyllum (whenever cladistic analyses are made with or without a backbone). Montsechia was an aquatic angiosperm living and reproducing below the surface of the water, similar to Ceratophyllum. Montsechia is Barremian in age, raising questions about the very early divergence of the Ceratophyllum clade compared with its position as sister to eudicots in many cladistic analyses. Lower Cretaceous aquatic angiosperms, such as Archaefructus and Montsechia, open the possibility that aquatic plants were locally common at a very early stage of angiosperm evolution and that aquatic habitats may have played a major role in the diversification of some early angiosperm lineages.When did early angiosperms begin to diversify ecologically? This question is currently unanswered. Age estimates of the divergence of crown-group angiosperms using molecular clock data vary considerably, although it is in the range of (max. 210–) often accepted, 150–140 (min. 130) million years (1–7). Parsimony reconstruction of early angiosperm habit suggests that they may have been shrubs living in “damp, dark, and disturbed” habitats (8). In contrast, many living aquatic angiosperms are basal in angiosperm phylogenies e.g., Nymphaeales in Amborella, Nymphaeales and Illiciales, Trimeniaceae-Austrobaileya (ANITA) or Ceratophyllales with the eudicots as commonly understood]. In the fossil record, we have found an aquatic angiosperm, Montsechia vidalii (Zeiller) Teixeira, which is an atypical plant fossil found in the Barremian (130–125 million years ago) freshwater limestone in the Pyrenees and Iberian Range in Spain. Montsechia (Fig. 1) lacks roots (no proximal or adventitious roots were found in more than 1,000 shoots examined) and shows flexible axes and two types of phyllotaxy and leaf morphology. The cuticle is very thin with rare stomata. The fruit is closed with a pore near the distal tip, indehiscent, and contains one unitegmic seed developed from an orthotropous and pendent ovule (Figs. 2 and ​and3).3). Cladistic analysis of these characters places Montsechia on the stem lineage basal to extant Ceratophyllum or a clade formed by Ceratophyllum and Chloranthaceae (Fig. 4) suggesting that mesangiosperms (non-ANITA angiosperms) existed 125 million years ago, as indicated by the tricolpate pollen record. Montsechia is well-adapted to a submerged aquatic habit. Montsechia is contemporaneous with another aquatic plant fossil, Archaefructus, indicating that some of the earliest angiosperms were fully aquatic very early in their ecological diversification.Open in a separate windowFig. 1.Long- and short-leaved forms of Montsechia vidalii. (A) The long-leaved specimen shows very flexuous branches and opposite, long leaves. LH02556. (Scale bar, 10 mm.) (B) The short-leaved specimen shows regularly developed lateral branches and tiny leaf rosettes. LH07198. (Scale bar, 10 mm.)Open in a separate windowFig. 2.Fruit and seed of Montsechia vidalii. The fruit shows a small apical pore (po). The funicle (f) of the single, upside-down seed (orthotropous pendent) is attached from the hilum (h) to the placenta (pl). (Scale bar, 500 µm.)Open in a separate windowFig. 3.Reconstructions of Montsechia vidalii. (A) The long-leaved form shows the opposite leaves and branches. (B) The short-leaved form shows the alternate phyllotaxy of leaves and branches bearing pairs of ascidiate, nonornamented fruits. (C and D) The fruit shows a small apical pore and a single seed developed from an orthotropous pendent ovule. The funicle arises from the placenta (near the micropyle) to the hilum (near the pollination pore). (C) Lateral view. (D) Front view. Diagram by O. Sanisidro, B.G., and V.D.-G.Open in a separate windowFig. 4.Most parsimonious position of Montsechia in a simplified tree derived from the matrix by Endress and Doyle (26) using the J & M backbone. Taxa in blue are considered ancestrally water-related (27). Diagram by C.C. and B.G.