Fossil Taiwannia from the Lower Cretaceous Yixian Formation of western Liaoning, Northeast China and its phytogeography significance

doi:10.3724/SP.J.1226.2018.00502

Abstract

Abstract:

Fossil Taiwania was discovered from the Lower Cretaceous Yixian Formation of Lingyuan City, western Liaoning Province, Northeast China. It is identified as a new species, Taiwania lingyuanensis sp. nov.. The present specimen is preserved as impressions with well defined leaf shoots system and reproductive structures. Leaves are dimorphic, spirally and imbricately arranged. They are scale-like on the main and cone-bearing branchlets, and subulate to falcate-subulate on the juvenile or sterile shoots. The seed cones are singly elliptic, ovate or elongate-ovate and terminally borne on ultimate shoots, bearing 22–24 scale-bracts complexes imbricately and helically arranged around the cone axis, the bracts are broad-ovate, rhomboidal or hexagonal with entire margins. Both the leafy shoots morphology and reproductive structures are similar to extant Taiwania. Furthermore, geological distribution and molecular biological evidences support that Taiwania is probably originated from the eastern Asia at least in the Early Cretaceous and widely distributed in the North Hemisphere thereafter.

Key words: Taiwannia, Lower Cretaceous, Yixian Formation, leaves, seed cones

MingZhen Zhang,BaoXia Du,PeiHong Jin,BaiNian Sun. Fossil Taiwannia from the Lower Cretaceous Yixian Formation of western Liaoning, Northeast China and its phytogeography significance[J].Sciences in Cold and Arid Regions, 2018, 10(6): 502-515.

Figures/Tables 8

Figure 1

Figure I

Figure II

Table 1

Morphological comparisons among the extant families of Coniferopsida"

Family		Leaf shape	Leaf arrangement	Seed or Seed cone type	Seed or seed cone arrangement	Seed or seed cone shape	Scale-bracts or bracts arrangement	Scale-bracts or bracts shape
Araucariaceae		Needle-like, broadly ovate or triangular-ovate, falcate, subulate, or triangular	Spirally or decussate	Seed cone	Solitary and terminal	Erect, ellipsoid, or ovoid to sub-globose	Spirally	Flabellate
Cupressaceae sensu lato	Cupressaceae sensu stricto	Scalelike or needle-like	Decussate or in whorls of 3	Seed cone	Solitary, terminal or axillary	Globose, ovoid, oblong or ellipsoid-cylindric	Decussate	Peltate
Taxodiaceae		Lanceolate, subulate, scale-like, or linear	Spirally scattered or decussate	Seed cone	Solitary, terminal or sub-terminal	Globose, ovoid, or cylindric-ovoid, ellipsoid	Spirally or decussate	Broadly ovate, triangular-ovate or triangular, peltate, rhombic, shield-shaped
Pinaceae		Scale-like, blade linear or needle-like	Spirally or cluster	Seed cone	Terminal or axillary	Cylindric or ovoid	Spirally and appressed	Peltate or flabellate
Sciadopityaceae		Scale-like, triangular or linear	Scattered or whorled	Seed cone	Terminal	Narrowly ovoid	Spirally	Broadly cuneate or fan-shaped
Cephalotaxaceae		Blade linear, linear-lanceolate, or lanceolate	Decussate or opposite, sessile or sub-sessile	Seed-bearing structure	Solitary	Obovate, ovoid or nearly globose with long pedunculate, drupelike	Decussate	Ovate
Taxaceae		Linear or lanceolate	Spirally arranged or decussate	Seed-bearing structure	Solitary or paired in axils of leaves	Ovoid or trigonous-ovoid, drupelike or nutlike, partially or completely enclosed in a succulent, saccate or cupular aril	At the base of the seed, overlapping or decussate	Triangular
Podocarpaceae		Blade scale-like, subulate, or linear to elliptic	Decussate, sub-opposite, or spirally arranged	Seed-bearing structure	Solitary, terminal or axillary	Drupelike or nutlike	Spirally	Apical bracts fertile, basal bracts often fused to form a receptacle

Table 1

Table 2

Morphological comparisons between taxodiaceous genera and the present specimen"

Genus	Leaves	Female cone shape and size	Bract-scale number, arrangement and shape	Ovules or Seed	References
Cunninghamia	Sessile, lanceolate or linear-lanceolate, spirally arranged, base decurrent, margin serrulate	1–3 together, terminal, globose, ovoid or cylindric-ovoid, 1.8–4.5 cm long and 1.2–4.0 cm wide	47–72, spirally arranged, sessile, connate proximally; tapered imbricate, broadly ovate or triangular-ovate, flat, large, base cordate, margin irregularly and finely serrulate	Ovules 1–3 per bract axil, Seeds 2–3	Fu et al., 1999c ; Farjon and Garcia, 2003; Schulz and Stützel, 2007; Ma et al., 2009
Taiwania	Sessile, falcate-subulate (juvenile stage) and scale-like (mature stage), spirally arranged, decurrent and imbricate	Shortly cylindric to ellipsoid, 1.0–2.2 cm long and 6–11 mm wide	20–25, tapered, cuneately narrowed into claw, appressed, with serrulate margins and an acute, slightly keeled apex	Ovules 1–2 per bract axil	Fu et al., 1999c ; Farjon and Garcia, 2003; Ma et al., 2009
Sequoia	Scale-like on main branches and linear on lateral branches, spirally arranged	Terminal, pendulous, pedunculate, ovoid-ellipsoid or ovoid, 0.9–2.5 cm long and 1.0–2.5 cm wide	21–28, shield-shaped, peltate with an apical depression, apically grooved, expanded into a rhomboid disc, occasionally with central mucro	Ovules 3–7 per bract axil, Seeds 2–5	Fu et al., 1999c ; Ma and Li, 2002; Farjon and Garcia, 2003; Ma et al., 2009
Sequoiadendron	Subulate and scale-like, spirally arranged	Terminal, pendulous, pedunculate, elliptic, 5–8 cm long and 3.0–5.5 cm wide	20–40, shield-shaped	Ovules 3–12 per bract axil	Fu et al., 1999c , Ma et al., 2009
Metasequoia	Linear, decussate	Ellipsoidal or spheroidal, 6.0–25 mm long and 6.0–25 mm wide	10–22, Shield-like, decussate, peltate with an apical depression 2.0–6.0 mm long and 5.0–17.0 mm wide	5–8 seeds in a row	Fu et al., 1999c ; Liu et al., 1999 ; Ma et al., 2009
Athrotaxis	Sessile, scale-like or needle-like, spirally arranged, loosely speading or closely appressed, with entire or serrate margins	Globose, 8–21 mm long	10–30, peltate, cuneate or peltate with tapered apex	3–6 seeds per complex	Miller and LaPasha, 1983; Farjon, 2005; Schulz et al., 2005 ; Eckenwalder, 2009; Ma et al., 2009
Cryptomeria	Spirally, crowded, linear or subulate, base decurrent	Globose or subglobose, ellipsoid, about 0.9–2.5 cm long and 1.0–2.5 cm wide	20–30, cuneate or peltate without an apical depression, 3–7 long tooth on the upper part of the seed-scale	2–5 seeds per complex	Fu et al., 1999c ; Ma et al., 2007 ; Ma et al., 2009
Glyptostrobus	Spirally, linear, linear-subulate and scale-like	Ellipsoid, 1.4–2.5 cm long and 0.9–1.5 cm wide	20–22, obovate, base cuneate, apical margin with 6–10 triangular, outwardly curved teeth adaxially.	Ovules 2 per bract axil	Fu et al., 1999c ; Ma et al., 2004
Taxodium	Linear or subulate spirally arranged	Globose or ellipsoid, 1.4–4.0 cm long and 1.3–3.0 cm wide	15–20, shield-shaped, peltate or rhombic, spirally arranged, not overlapping, apex irregularly quadrangular, with longitudinal rib on the upside margins	Ovules 2 per bract axil, Seeds 2 per cone scale	Fu et al., 1999c ; Kunzmann et al., 2009 ; Ma et al., 2009
Taiwania lingyuanensis	Subulate or falcate-subulate on the young branches or sterile branches, scale-like on the main branch and fertile branches	Ellipsoid or cylindric, 10.7–13.0 mm long and 6.5–7.5 mm wide	22–24, spirally arranged, broad-ovate, rhomboidal or hexagon, base broad-rhomboidal, triangular or round and apex gradually tapering	Ovules 2–3	The present paper

Table 2

Figure 2

Table 3

Morphological comparisons among the reliable fossil records of Taiwania and Taiwania-like species "

Species	Fossil type	Age	Locality	Leaves	Seed cone	References
T. eocenica	Seed cone	Eocene	Japan	–	12 mm long and 8 mm wide	Matsuo, 1967
T. fushunensis	Leafy shoot bearing seed cone	Eocene	China	Scale-like	19 mm long and 14 mm wide	Endo, 1942
T. paracryptomeroides	Seed cones	Miocene	Germany	–	15 mm long and 6 mm wide	Kilpper, 1968
T. cf. paracryptomeroides	Leaves	Miocene	Germany	Subulate, 3–5 mm long and 1.5–2.0 mm wide	–	Mai and Walther, 1991
T. mesocryptomeroides	Leafy shoot bearing seed cone	Late Cretaceous (Maastrichtian)	Japan	Subulate and scale-like	6–8 mm long and 4–5 mm wide	Matsuo, 1970
T. japonica	Leafy shoot bearing seed cone	Miocene and Pilocene	Japan	Subulate and scale-like, subulate leaves 3–14 mm long and 1–2 mm wide	10 mm long and 5 mm wide	Ishida, 1970
T. cretacea	Seed cones and leaves	Cretaceous	Russia	Subulate, 7 mm long and 2.5 mm wide	17 mm long and 15 mm wide	Samylina, 1988
T. microphylla	Leaves	Late Cretaceous (Cenomanian; Turonian)	New Siberia Island	Subulate leaves about 2–7 mm long and 0.7–2.5 mm wide, scale leaves about 1.25 mm long and 1 mm wide	–	Sveshnikova and Budantsev, 1969
T. rarytkiniana	Leaves	Upper Eocene	Russia	Subulate leaves about 6–7 mm long and 3.5–4.0 mm wide	–	Akhmetiev and Samsonenko, 1997
T. schaeferi	Seed cone and leaves	Paleocene and Miocene	Germany	Subulate leaves about 7 mm long and 3 mm wide, scale leaves 2.5–4.0 mm long and 1.5–2.0 mm wide	7–8 mm long and 3–5 mm wide	Schloemer-J?ger, 1958; Kilpper, 1968; J?hnichen, 1998
T. cf. schaeferi	Leaves	Oligocene	Germany	Subulate, about 12–14 long and 2 mm wide	–	Walther, 1999
Parataiwania nihongii	Seed cone	Late Cretaceous (Coniacian-Santonian)	Hokkaido, Japan	–	Elliptic, 22 mm long and 16 mm wide, 3 ovules per cone scale	Nishida et al., 1992
Mikasastrohus hokkaidoensis	Seed cone	Late Cretaceous (Coniacian-Sanonian)	Hokkaido, Japan	Helically arranged, rhombic in transverse section	Ovoid, 3–4 cm long and up to 2.0–3.5 cm in diameter	Saiki and Kimura, 1993
T. lingyuanensis	Leafy shoot bearing seed cones	Early Cretaceous (Aptian)	China	Subulate leaves 1.5–4.0 mm long and 0.7–1.2 mm wide, scale leaves 2–3 mm long and 0.8–2.5 mm wide	Elliptic, ovate or elongate-ovate, 10.7–13 mm long and 6.5–7.5 mm wide	The present paper

Table 3

Figure 3

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