Sciences in Cold and Arid Regions ›› 2018, Vol. 10 ›› Issue (6): 502-515.doi: 10.3724/SP.J.1226.2018.00502

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

MingZhen Zhang1,*(),BaoXia Du2,PeiHong Jin1,2,BaiNian Sun2   

  1. 1 Key Laboratory of Petroleum Resources, Gansu Province/Key Laboratory of Petroleum Resources Research, Institute of Geology and Geophysics, Chinese Academy of Sciences, Lanzhou, Gansu 730000, China
    2 Key Laboratory of Mineral Resources in Western China (Gansu Province) and School of Earth Science, Lanzhou University, Lanzhou, Gansu 730000, China
  • Received:2018-10-19 Accepted:2018-11-16 Online:2018-12-01 Published:2018-12-29
  • Contact: MingZhen Zhang
  • Supported by:
    We are grateful to Prof. Xiaolin Wang from the Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China for providing the present fossil specimen. This work was conducted under the 973 Program of China (No. 2012CB822003), the National Natural Science Foundation of China (No. 41402007, 41602023) and the Fundamental Research Funds for the Central Universities (Lzujbky-2014-132).


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

Figure 1

Location of the fossil site in western Liaoning, Northeast China"

Figure I

Leafy shoots morphology of Taiwania lingyuanensis (1. Fossil specimen of Taiwania lingyuanensis, scale bar=20 mm; 2. Leafy shoots and seed cones of Taiwania lingyuanensis, scale bar=10 mm; 3. Lateral sterile and fertile branches of Taiwania lingyuanensis, scale bar=5 mm; 4. main branch of Taiwania lingyuanensis, scale bar=5 mm; 5–7. The main and lateral branches of Taiwania lingyuanensis, scale bar=5 mm; 8. Leaf morphology of the main branches, scale bar=2 mm; 9–10. Leaf morphology of the lateral sterile branches, scale bar=2 mm) "

Figure II

Seed cone morphology of Taiwania lingyuanensis (1–3. singly and terminally inserted seed cones, scale bar=5 mm; 4. scale-like leaves and rhomboidal scale-bracts of Taiwania lingyuanensis, shows the distinctive keel on the abaxial surface, scale bar=5 mm; 5. spirally and imbricate scale-bracts of Taiwania lingyuanensis, shows the trace of seeds, scale bar=5 mm; 6. scale-bracts morphology of Taiwania lingyuanensis, shows the trace of the small lob with irregularly sinuate margins, scale bar=5 mm; 7. scale-bracts morphology at the base of the seed cone, scale bar=2 mm; 8–9. scale-bracts morphology at the middle and upper part of the seed cone, scale bar=2 mm) "

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 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

Figure 2

Comparison of morphological structures between the holotype and the extant Taiwania (a: foliage of the holotype; b: foliage of Taiwania cryptmerioides), derive from Delectis Florae Reipublicae Popularis Sinicae Agendae Academiae Sinicae, 1978 "

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

Figure 3

Fossil and extant distribution of Taiwania (Data from: Endo, 1942; Schloemer-J?ger, 1958; Matsuo, 1967; Kilpper, 1968; Sveshnikova and Budantsev, 1969; Ishida, 1970; Matsuo, 1970; Samylina, 1988; Mai and walther, 1991; Nishida et al., 1992 ; Akhmetiev and Samsonenko, 1997; J?hnichen, 1998; McIver and Basinger, 1999; Walther, 1999; LePage, 2009) "

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