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On Montsechia, an Angiospermoid Plant from the Lower Cretaceous of Las Hoyas, Spain: New Data and Interpretations

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On Montsechia, an Angiospermoid Plant from the Lower Cretaceous of Las Hoyas, Spain: New Data and Interpretations Acta Palaeobotanica 51(2): 181–205, 2011 On Montsechia, an angiospermoid plant from the Lower Cretaceous of Las Hoyas, Spain: new data and interpretations VALENTIN KRASSILOV Institute of Evolution, University of Haifa, Israel; e-mail: [email protected] Received 13 June 2011; accepted for publication 20 October 2011 ABSTRACT. Montsechia from the Las Hoyas Locality, Cuenca, Spain, previously described as a fl oating plant with whorled leaves is re-interpreted as having a dimorphic long shoot – short shoot branching system with heterophyllous elongate and scaly leaves. The stomata and trichomes are observed in the scaly leaves. The growth habit is interpreted as heloxerophytic over the coastal marshland habitats. The ovulate cupules are terminal on the reproductive short shoots and contain a solitary ovule. Pollen grains are occasionally found on the nucellus betraying a gymnospermous pollination mode. The pollen tube morphology is like in gnetophytes and angiosperms. Some seed-like structures on scale leaves and bud cataphylls contain larval remains and are interpreted as nematode galls. The taxonomic affi nities of Montsechia are with the helophytic proangiosperms, in particular the Baisiales. The Montsechia – Montsechites (“Ranunculus”) assemblage of Montsec and La Hoyas is comparable to the mid-Cretaceous proangiosperm assemblages of Transbaikalia, Mongolia and eastern China. KEYWORDS: Montsechia, early angiosperm, proangiosperms, nematode galls, Cretaceous, Las Hoyas locality, Spain INTRODUCTION The pre-Albian angiosperm records are of great has remained controversial ever since. This interest bearing on the problem of angiosperm paper reports on my further results concern- origin and the rates of initial angiosperm evolu- ing growth form and reproductive structures, tion. Montsechia vidalii (Zeiller 1902) Teixeira as well as the seed-like bodies interpreted as 1954 from the supposed Barremian deposits of nematode galls. As in the present day wheat Spain ranks among the earliest angiosperm- nematodes, the galls seem to have been quite like fossils supposedly indicating a long pre- similar to the seeds. A designation “seed-like Albian history of the group (Daviero-Gomez body” is here applied to uncertain cases. et al. 2006, Gomez et al. 2006). However, since My results based on a limited material angiosperm characters are reported in diverse must be considered as preliminary step toward non-angiospermous (proangiospermous) Creta- understanding Montsechia and its signifi cance ceous and even pre-Cretaceous plants (reviewed for the problem of angiosperm origins. in Krassilov 1997, 2009), a cautious revision of such fi nds is necessary before evolutionary implications are discussed. MATERIAL AND METHODS Both the interpretation of Montsechia as angiosperm (Gomez et al. 2006) and my pre- Montsechia is abundant in laminated carbonates liminary conclusion on gnetophytic affi nities of at Las Hoyas outcrop of Iberian Basin, from which I obtained 30 specimens courtesy of Alexander Ban- this plant (Krassilov 2006) were presented at nikov and Tatyana Kodrul who visited this locality the 7th European Paleobotanical and Palyno- during post-congressional fi eld trips in 2005 and 2008. logical Conference, Prague, 2006, and the issue A larger slab about 20 × 15 cm presents several shoot 182 fragments and several dozens detached scale leaves and bract compressions bearing seed-like bodies The material was studied on the rock surface as well as on fi lm transfers and in bulk maceration residues. Altogether, 70 seed-like bodies were cleared for light microscopy and mounted for SEM. The microphoto- graphs were obtained with the stereomicroscope Leica 300, dissecting light microscope Nikon Eclipse and scanning electron microscope FE1 Quanta 200. The material is stored in the palaeobotanical depository of the Institute of Evolution, University of Haifa, No. SLH 1 – 70. VEGETATIVE MORPHOLOGY The traditional interpretation of Montsechia as having successive leaf whorls on monomor- phic shoot axes is incorrect. Gomez et al. (2006) have discerned two growth forms, with long decussate leaves and scaly spiral leaves, but failed to recognize dimorphic shoot systems and heterophylly in both the long-leafed and short-leafed varieties. The leaves are decus- sate rather than whorled, the apparent whorls consisting of adjacent leafy short shoots at the Fig.1. Montsechia vidalii (Zeiler) Teixeira, long-leafed last order branching nodes. branching shoot The material at hand represents leafy shoots of three orders of branching. The fi rst The scaly form is also heterophyllous (Fig. 2), order axes about 1.5 mm thick give off disti- but the elongate leaves are mostly shed leaving chous lateral branches that are opposite at the raised leaf cushions that protrude between the regularly spaced branching nodes. The lateral branches are slender, upcurved, of unequal length, with decussate leaves, subtending axil- lar short shoots that can be missing from the axils of a few apical leaf pairs (Pl. 1, fi g. A). Both long and short shoots are heterophyl- lous, foliated with elongate and scaly leaves. One or another leaf shape prevails distinguish- ing the “leafy” and “scaly” forms (Pl. 1, fi gs A–C). In the leafy form (Fig. 1), the prevailing leaf shapes are linear, falcate, sessile, bluntly pointed, thick and apparently succulent, with strong fi bers transpiring through the epider- mis (Pl. 2, fi gs A–C; Pl. 3, fi g. B). The leaf apices are minutely serrate with microscopic teeth (Pl. 3, fi g. B). Their axillar short shoots are distichous, opposite at the branching nodes, developing as small knobs and varying from hemispherical to club-shaped when fully developed. The short shoot leaves are hetero- morphic, with a few scale leaves at the base followed by a tuft of overlapping transitional and elongate leaves. Two intermingling leaf Fig. 2. Montsechia vidalii (Zeiler) Teixeira, branching scale- tufts of the opposite short shoots produce an leafed shoot with two buds (arrows)and the buds magnifi ed; apparent “leaf whorl” of the node. ax, shoot axis sprouting from between the cataphylls 183 scales. A few welting leaves may persist (Pl. 1, sporogonia). In both the leafy and scaly varie- fi g. B). The short shoots vary from hemispheri- ties, these structures are borne terminally on cal to shortly cylindrical, uniformly covered the short shoots developing in the leaf axils of with scale leaves that are shortly triangular, penultimate branches, but in the leafy variety acute, imbricate, decurrent, with the free part they are solitary and scattered (Pl. 4, fi g. B), slightly spreading. whereas as they are more frequent and aggre- The scale leaf cuticle is moderately thick, gated on reproductive shoots. In Fig. 2, the papillate, with a fringe of transversely elon- scale-leafed branching system bears two coni- gate cells and with a few stomata in the cen- cal buds (arrows) with the shoot axis sprout- tral part of their adaxial face (Pl. 3, fi gs A, B). ing between the overlapping cataphylls. A few The stomata are irregularly scattered on the cataphylls persist on the axis of the sprout- densely trichomate leaf surface, with sunken ing long shoots ( Pl. 5, fi gs C, D) that produce guard cells and irregular subsidiary cells that several pairs of fertile short shoots (Pl. 5, fi gs are smaller than the pavement cells. A, B). A leafy shoot in Pl. 4 fi g. B produces dis- tichous short shoots only one of which is fer- tile (arrow). The opposite short shoot bears a terminal tuft of elongate leaves topologically equivalent to the ovulate structure the exter- nal coat of which is formed of incompletely connate leaves adhering to the inner body. In the scale-leafed reproductive shoot system, the fertile short shoots produce one to several pairs of small falcate leaves before the ter- minal ovulate structure. Slender scale leaves spread sideways from the elongate terminal body (Pl. 5, fi g. B) that is notched at the apex and shows a median split between the constit- uent bracts. This structure was left intact for documentation, but an analogous if but acu- minate and slightly falcate structure in Pl. 5, fi g. A was macerated yielding a cutinized inner Fig. 3. Montsechia vidalii (Zeiler) Teixeira conducting tissue body Pl. 5, fi g. E). of a shoot axis (Pl. 2, fi g. C), arrows on the bundles of spiral These observations seem congruous with tracheids or tracheid fi bers surrounded with metatracheal interpretation of the external seed coat as parenchyma a cupule formed of bracteate appendages, A few anatomical features are discernible with the foliage leaves down the axis partly in cleared shoot axes. The conducting tissue involved in fusion. Their shapes vary from is organized in a few parallel bundles showing ovate, shortly apiculate, broadly truncate at spiral tracheids or tracheary fi bers surrounded base to elongate-elliptic, cuneate or slightly with tabloid cells of metatracheal parenchyma cordate at base, apically acuminate or pungent, (Fig. 3). about.1.5–3.0 mm long. The cupule walls are thick compressions with indistinct longitudinal folds or ribs, fi brous, showing longitudinal cell REPRODUCTIVE STRUCTURES fi les. Irregular pits on the walls apparently rep- resent hair bases. They contain a solitary ovule Attached to Montsechia shoots are the that is epitropous (curved up toward the apex), bladder-like thick walled, but feebly cuti- elongate or elliptic, straight or slightly curved, nized structures investing a solitary inner keeled, about 1.0–1.5 mm long.. The chalazal body which is well cutinized and detachable end is smoothly rounded; the micropylar end is by maceration, but actually never found dis- bluntly pointed or shortly mucronate. persed (therefore not assignable to Spermatites Cleared ovules show a slender inner integu- Miner 1935, a form genus for dispersed seed- ment one cell layer thick, extending up to 2/3 like bodies probably comprising also bryophyte of the nucellus length (Pl. 6, fi g. C; Pl. 7, fi g. C), 184 rest shows a tube 5 μm wide emerging from between the narrow saccus lobes and turn- ing down the nucellus (Pl.
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