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ephedroids from the Early Cretaceous Yixian Formation in Liaoning ...

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246 C. Ryd<strong>in</strong> et al.: <strong>Cretaceous</strong> <strong>ephedroids</strong> <strong>from</strong> Ch<strong>in</strong>a<br />

rounded, elliptic, 2.5–3.5 mm long, 1.2–2 mm<br />

wide.<br />

Etymology. From <strong>the</strong> robust morphology<br />

of <strong>the</strong> plant.<br />

Holotype. PB20719. (Figs. 16, 18).<br />

Paratype. (Counterpart) PB20720. (Fig. 17).<br />

Locality. Huangbanjigou Section, Beipiao,<br />

Liaon<strong>in</strong>g, Ch<strong>in</strong>a.<br />

Stratigraphic position and age. Jianshangou<br />

Bed, <strong>Yixian</strong> <strong>Formation</strong>, <strong>Early</strong> <strong>Cretaceous</strong><br />

(Barremian-early Aptian?).<br />

Description and comments. The type material<br />

of Liaoxia robusta comprises two compression<br />

fossils of an ephedroid, female plant,<br />

part and counterpart, with oxidized organic<br />

material preserved. The shoot is 17 cm long<br />

(Fig. 16), but on <strong>the</strong> counterpart, only <strong>the</strong><br />

uppermost 8 cm are preserved (Fig. 17). No<br />

<strong>in</strong>ternal or external anatomical details are<br />

preserved. Precipitations surround <strong>the</strong> plant<br />

rema<strong>in</strong>s and make stems and leaves appear<br />

wider than <strong>the</strong>y are.<br />

Stems are erect with nodes and <strong>in</strong>ternodes,<br />

1.5–2 mm thick, thicker at nodes, and have<br />

longitud<strong>in</strong>al striations-ridges. Phyllotaxis is<br />

opposite and decussate with multiple axillary<br />

units. Leaves are l<strong>in</strong>ear, 15–20 mm long and<br />

appear 1.5–2 mm wide, but <strong>the</strong> width also<br />

comprises precipitates and <strong>the</strong> leaves were<br />

probably only 0.5–1 mm wide. Indist<strong>in</strong>ct<br />

rema<strong>in</strong>s of three or four ve<strong>in</strong>s are present on<br />

some leaves. The plant has about 15 ovulate<br />

cones that are rounded to ellipsoid <strong>in</strong> longitud<strong>in</strong>al<br />

outl<strong>in</strong>e. Cones vary <strong>in</strong> length between 7<br />

and 16 mm, but are all about 6–8 mm wide,<br />

sessile or pedunculate, and are positioned<br />

axillary at nodes or term<strong>in</strong>ally on branches.<br />

Cones comprise between approximately 4 and<br />

10 pairs of reflexed and decussately arranged<br />

bracts. The bracts are ovate, 4–7 mm long,<br />

with a long and acute to attenuate apex. Seeds<br />

are rounded to elliptic, 2.5–3.5 mm long and<br />

1.2–2 mm wide, but no cell structures or o<strong>the</strong>r<br />

anatomical details are preserved. There is one<br />

seed <strong>in</strong> <strong>the</strong> axil of all cone bracts, <strong>in</strong>clud<strong>in</strong>g<br />

proximal bracts.<br />

This plant differs <strong>from</strong> o<strong>the</strong>r species of<br />

Liaoxia <strong>in</strong> that all features are larger. Fur<strong>the</strong>r,<br />

it has multiple axillary branch<strong>in</strong>g, which has<br />

o<strong>the</strong>rwise only been observed for Liaoxia<br />

elongata and L. longibractea, and <strong>the</strong>se species<br />

are clearly different <strong>from</strong> L. robusta <strong>in</strong> <strong>the</strong><br />

characters of <strong>the</strong> cones. Liaoxia robusta differs<br />

<strong>from</strong> Ephedra <strong>in</strong> <strong>the</strong> presence of seeds <strong>in</strong> <strong>the</strong><br />

axil of all bracts, not only <strong>in</strong> distal pairs as <strong>in</strong><br />

extant Ephedra.<br />

A detached w<strong>in</strong>ged seed occurs <strong>in</strong> close<br />

association with Liaoxia robusta (Fig. 18).<br />

The seed body of this isolated seed is of<br />

approximately <strong>the</strong> same length as <strong>the</strong> <strong>in</strong> situ<br />

seeds and <strong>the</strong> possibility that it orig<strong>in</strong>ates<br />

<strong>from</strong> a Liaoxia cone cannot be ruled out, but<br />

<strong>the</strong> shape of <strong>the</strong> seed and <strong>the</strong> nature of <strong>the</strong><br />

w<strong>in</strong>g suggest that it was more likely produced<br />

by one of <strong>the</strong> conifers reported for <strong>the</strong> <strong>Yixian</strong><br />

<strong>Formation</strong>.<br />

Liaoxia longibractea Ryd<strong>in</strong>, S.Q. Wu et Friis<br />

sp. nov.<br />

Specific diagnosis. As for <strong>the</strong> genus with <strong>the</strong><br />

follow<strong>in</strong>g dist<strong>in</strong>ctions. Branches c. 1.5 mm<br />

wide, with multiple axillary branch<strong>in</strong>g. Cones<br />

sessile to very shortly pedunculate, rounded,<br />

3.5–4 mm long ()15 mm <strong>in</strong>clud<strong>in</strong>g bract<br />

apex), c. 2–3 mm wide, with 1–2 pairs of<br />

bracts. Cone bracts reflexed, 10–15 mm long,<br />

extended with a setose apex, two parallel<br />

ve<strong>in</strong>s. Seeds elliptic, 2.5–4.0 mm long, c.<br />

1 mm wide.<br />

Etymology. Named after <strong>the</strong> very long and<br />

narrowly extended cone bracts.<br />

Holotype. PB20725 (Fig. 19).<br />

Paratype. PB20726 (<strong>the</strong> counterpart,<br />

Fig. 20).<br />

Locality. Fanzhangzi Section, L<strong>in</strong>gyuan,<br />

Liaon<strong>in</strong>g, Ch<strong>in</strong>a.<br />

Stratigraphic position and age. Dawangzhangzi<br />

Bed, <strong>Yixian</strong> <strong>Formation</strong>, <strong>Early</strong><br />

<strong>Cretaceous</strong> (Barremian-early Aptian?).<br />

Description and comments. The type material<br />

of Liaoxia longibractea comprises two<br />

relatively well-preserved compression fossils<br />

(part and counterpart) with small amounts of<br />

oxidized organic material preserved. No <strong>in</strong>ternal<br />

or external anatomical details are pre-

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