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

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

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244 C. Rydin et al.: Cretaceous ephedroids from China about 1.7–2.7 mm wide. Cones are sessile and placed axillary at nodes or terminally on branches. Each cone has between 6 and 12 pairs of erect to slightly reflexed bracts that appear decussately arranged (Fig. 10). Bracts are narrow and ovate, 1.5–2.0 mm long, with an acute apex. Seeds or microsporangiate organs have not been observed. This fossil is similar to Liaoxia chenii in vegetative morphology, but the cones are more elongate in shape. Liaoxia acutiformis is further distinguished from L. chenii by having a higher number of bracts, probably up to 12 pairs, in each cone, in contrast to L. chenii that has 2–6 pairs in each cone. The apex of the cone bracts differs in being acute to obtuse in L. acutiformis while it is acute to attenuate in L. chenii. Liaoxia elongata Rydin, S.Q. Wu et Friis sp. nov. Specific diagnosis. As for the genus with the following distinctions. Branches 0.4–1.5 mm thick, sometimes with multiple axillary nodes. Leaves absent. Cones narrowly elongate, 5– 10 mm long, c. 2–3 mm wide, with 8–12 pairs of bracts. Cone bracts erect, 2.5–4 mm long, with an attenuate apex. Etymology. Named after the narrowly elongated cones. Holotype. PB20724 (Figs. 14–15). Paratype. PB20723 (Figs. 12–13). Locality. Fanzhangzi Section, Lingyuan, Liaoning, China. Stratigraphic position and age. Dawangzhangzi Bed, Yixian Formation, Early Cretaceous (Barremian-early Aptian?). Description and comments. The holotype of Liaoxia elongata comprises the uppermost 5.5 cm of the plant (Figs. 14–15). An additional specimen is a shoot with slightly larger structures, c. 4 cm long and 4 cm wide, repre- senting the central part of the plant (Figs. 12– 13). No anatomical details are preserved. The stems are erect with nodes and internodes. Branching is opposite and the paratype has multiple axillary units (Fig. 12). Stems are 0.4–1.5 mm thick, thicker towards the base and at nodes, and have very weak remnants of longitudinal striations. Leaves are absent. Cones are axillary at nodes or terminally on branches and may be sessile or pedunculate. They are narrowly elongate (Figs. 13–14) and vary in length between 5–6 mm in the holotype to 6–10 mm in the paratype, but are all 2–3 mm wide. One cone on the paratype is smaller and rounded, c. 3 · 3 mm (arrow in Fig. 12). Each cone comprises 8–12 pairs of erect and decussately arranged bracts. The bracts are probably very narrowly ovate, c. 2.5–4 mm long, with attenuate apex. Seeds or microsporangiate organs have not been observed. These plants are similar to Liaoxia changii (Cao et al. 1998) in the over all ephedroid habit, and the leaf-less stems, but differ from Liaoxia changii mainly in that they have cones with more bracts, which are erect and have a distinctly pointed, attenuate apex. The cone bracts of Liaoxia changii are slightly reflexed and have a less distinctly pointed, acute apex. Liaoxia robusta Rydin, S.Q. Wu et Friis sp. nov. Specific diagnosis. As for the genus with the following distinctions. Branches 1.5–2 mm thick, with multiple axillary units. Leaves at least 15–20 mm long, 0.5–1.0 mm wide. Cones obovate to elongate 7–16 mm long, c. 6–8 mm wide, with 4–10 pairs of bracts. Cone bracts reflexed, 4–7 mm long, ovate with acute to attenuate apex, one-two parallel veins. Seeds c Figs. 1–4. Liaoxia chenii Cao et S.Q. Wu (emend. Rydin, S.Q. Wu et Friis). 1 The holotype of Liaoxia chenii (PB17800). 2 New specimen (PB20718); the detached apical part of a reproductive shoot. 3–4 New specimen (PB20717); the uppermost part of a branched plant. Note the seed and the thin leaf indicated by arrows in Fig. 3. Scale bars = 1 cm in Figs. 1, 2, 4; 0.3 cm in Fig. 3. S = seed; L =leaf

C. Rydin et al.: Cretaceous ephedroids from China 245

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

about 1.7–2.7 mm wide. Cones are sessile and<br />

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

branches. Each cone has between 6 and 12<br />

pairs of erect to slightly reflexed bracts that<br />

appear decussately arranged (Fig. 10). Bracts<br />

are narrow and ovate, 1.5–2.0 mm long, with<br />

an acute apex. Seeds or microsporangiate<br />

organs have not been observed.<br />

This fossil is similar to Liaoxia chenii <strong>in</strong><br />

vegetative morphology, but <strong>the</strong> cones are more<br />

elongate <strong>in</strong> shape. Liaoxia acutiformis is fur<strong>the</strong>r<br />

dist<strong>in</strong>guished <strong>from</strong> L. chenii by hav<strong>in</strong>g a higher<br />

number of bracts, probably up to 12 pairs, <strong>in</strong><br />

each cone, <strong>in</strong> contrast to L. chenii that has 2–6<br />

pairs <strong>in</strong> each cone. The apex of <strong>the</strong> cone bracts<br />

differs <strong>in</strong> be<strong>in</strong>g acute to obtuse <strong>in</strong> L. acutiformis<br />

while it is acute to attenuate <strong>in</strong> L. chenii.<br />

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

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 0.4–1.5 mm<br />

thick, sometimes with multiple axillary nodes.<br />

Leaves absent. Cones narrowly elongate, 5–<br />

10 mm long, c. 2–3 mm wide, with 8–12 pairs<br />

of bracts. Cone bracts erect, 2.5–4 mm long,<br />

with an attenuate apex.<br />

Etymology. Named after <strong>the</strong> narrowly<br />

elongated cones.<br />

Holotype. PB20724 (Figs. 14–15).<br />

Paratype. PB20723 (Figs. 12–13).<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 holotype of<br />

Liaoxia elongata comprises <strong>the</strong> uppermost<br />

5.5 cm of <strong>the</strong> plant (Figs. 14–15). An additional<br />

specimen is a shoot with slightly larger<br />

structures, c. 4 cm long and 4 cm wide, repre-<br />

sent<strong>in</strong>g <strong>the</strong> central part of <strong>the</strong> plant (Figs. 12–<br />

13). No anatomical details are preserved.<br />

The stems are erect with nodes and <strong>in</strong>ternodes.<br />

Branch<strong>in</strong>g is opposite and <strong>the</strong> paratype<br />

has multiple axillary units (Fig. 12). Stems are<br />

0.4–1.5 mm thick, thicker towards <strong>the</strong> base<br />

and at nodes, and have very weak remnants of<br />

longitud<strong>in</strong>al striations. Leaves are absent.<br />

Cones are axillary at nodes or term<strong>in</strong>ally on<br />

branches and may be sessile or pedunculate.<br />

They are narrowly elongate (Figs. 13–14) and<br />

vary <strong>in</strong> length between 5–6 mm <strong>in</strong> <strong>the</strong> holotype<br />

to 6–10 mm <strong>in</strong> <strong>the</strong> paratype, but are all<br />

2–3 mm wide. One cone on <strong>the</strong> paratype is<br />

smaller and rounded, c. 3 · 3 mm (arrow <strong>in</strong><br />

Fig. 12). Each cone comprises 8–12 pairs of<br />

erect and decussately arranged bracts. The<br />

bracts are probably very narrowly ovate, c.<br />

2.5–4 mm long, with attenuate apex. Seeds or<br />

microsporangiate organs have not been<br />

observed.<br />

These plants are similar to Liaoxia changii<br />

(Cao et al. 1998) <strong>in</strong> <strong>the</strong> over all ephedroid<br />

habit, and <strong>the</strong> leaf-less stems, but differ <strong>from</strong><br />

Liaoxia changii ma<strong>in</strong>ly <strong>in</strong> that <strong>the</strong>y have<br />

cones with more bracts, which are erect and<br />

have a dist<strong>in</strong>ctly po<strong>in</strong>ted, attenuate apex. The<br />

cone bracts of Liaoxia changii are slightly<br />

reflexed and have a less dist<strong>in</strong>ctly po<strong>in</strong>ted,<br />

acute apex.<br />

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

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 1.5–2 mm<br />

thick, with multiple axillary units. Leaves at<br />

least 15–20 mm long, 0.5–1.0 mm wide. Cones<br />

obovate to elongate 7–16 mm long, c. 6–8 mm<br />

wide, with 4–10 pairs of bracts. Cone bracts<br />

reflexed, 4–7 mm long, ovate with acute to<br />

attenuate apex, one-two parallel ve<strong>in</strong>s. Seeds<br />

c<br />

Figs. 1–4. Liaoxia chenii Cao et S.Q. Wu (emend. Ryd<strong>in</strong>, S.Q. Wu et Friis). 1 The holotype of Liaoxia chenii<br />

(PB17800). 2 New specimen (PB20718); <strong>the</strong> detached apical part of a reproductive shoot. 3–4 New specimen<br />

(PB20717); <strong>the</strong> uppermost part of a branched plant. Note <strong>the</strong> seed and <strong>the</strong> th<strong>in</strong> leaf <strong>in</strong>dicated by arrows <strong>in</strong><br />

Fig. 3. Scale bars = 1 cm <strong>in</strong> Figs. 1, 2, 4; 0.3 cm <strong>in</strong> Fig. 3. S = seed; L =leaf

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