A New Species of the Extinct Genus Austrohamia (Cupressaceae

Journal of Systematics and Evolution 50 (1): 72–82 (2012) doi: 10.1111/j.1759-6831.2011.00165.x

Research Article A new species of the extinct genus Austrohamia (Cupressaceae s.l.)in the Daohugou Jurassic ﬂora of China and its phytogeographical implications 1Jian-Wei ZHANG 2Ashalata D’ROZARIO 3Li-Jun WANG 4Ya L I 1Jian-Xin YAO∗ 1(Institute of Geology, Chinese Academy of Geological Sciences, Beijing 100037, China) 2(Department of Botany, Narasinha Dutt College, Howrah 711101, India) 3(College of Ethnology and Sociology, South Central University for Nationalities, Wuhan 430074, China) 4(State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China)

Abstract A new conifer, Austrohamia acanthobractea, sp. nov., is described from the Jurassic Daohugou flora, Inner Mongolia Autonomous Region, China. The material consists of impressions represented by well-preserved leafy twigs and branches as well as ovulate cones. Leafy shoots with at least two orders of branching; ultimate branchlets alternate or sub-opposite with helically arranged leaves, decurrent at base with distal rounded tip; dorsiventrally flattened and univeined. Ovuliferous cones elliptical, less than 1 cm long, terminally borne on ultimate and penultimate branches, composed of helically arranged bracts with ovules disposed on their adaxial surfaces. The presence of similar, if not identical taxa, on both sides of the Pacific indicates the cosmopolitan distribution of primitive Cupressaceae between East Asia (Eurasia) and South America in the Pangaea. Key words Austrohamia acanthobractea, China, Cupressaceae s.l., Jurassic, Pangaea, phytogeography.

Lacustrine deposits of tuffaceous mudstone, shale, 1994; Stefanovic et al., 1998; Kusumi et al., 2000), and and siltstone of the Daohugou area (Ren et al., 2002; morphological features (Hart, 1987; Miller, 1988; Far- Liu et al., 2004, 2010; Wang et al., 2005) in the In- jon et al., 2002), combined datasets (Gadek et al., 2000), ner Mongolia Autonomous Region, China (Fig. 1), comparative immunological analyses (Price & Lowest- which border the “Jehol Biota” in the western Liaon- ein, 1989), and development of the ovule and ovulate ing Province, China, are rich in well-preserved fossils cones (Farjon & Ortiz Garc´ıa, 2003), indicate that the of animals, such as freshwater conchostracans, bivalves, traditional Taxodiaceae is not a monophyletic group (is insects, caudata, pterosaurs, mammals, agnathans, and paraphyletic). The separation of Cupressaceae s.s. and other fauna (Zhang, 2002; Ji, 2007; Evans & Wang, Taxodiaceae showed no differences in any consistent 2009; Liu et al., 2010) and plants, such as Ginkgopsida, characteristics, and it suggests that both families should Coniferopsida, Filicopsida, and Cycadopsida (Zhang, be combined under the earlier name Cupressaceae (Cu- 2002; Ren & Yin, 2003; Ji et al., 2005; Jiang, 2006; pressaceae s.l.). Zhang et al., 2006, 2011; Zhou et al., 2007). The de- Based on the plesiomorphic characteristics posit is part of the Middle Jurassic to Late Jurassic (Brunsfeld et al., 1994; Gadek et al., 2000; Kusumi “Yanliao Biota” (Hong, 2009; Liu et al., 2010). Among et al., 2000; Farjon et al., 2002; Farjon, 2005), that is, the Coniferopsida, four genera have been described or pollen cones occurring in clusters or in aggregations, identiﬁed: Schizolepis, Pityospermum, Pityocladus, and pollen sacs in numbers of 4–5, absence of papillae on Elatocladus (Yanliaoia?), and only the last one is related cone scales, foliate nature of mature cone scale, domi- to the family Cupressaceae (Zhang, 2002; Ji et al., 2005; nance of bract scale of the cone, and cone scale fused Zhang et al., 2011). with coriaceous ovuliferous cone at maturity, the genera Studies based on their phenetic (Eckenwalder, previously placed in the “Taxodiaceae” are in the basal 1976), molecular (Chase et al., 1993; Brunsfeld et al., position of the family. Under the Cupressaceae s.l., some fossil genera, such as Elatides (Heer) Harris, Sewardiodendron (Florin) Yao, Zhou & Zhang, and Austrohamia Escapa, Received: 15 April 2011 Accepted: 24 August 2011 ∗ Author for correspondence. E-mail: [email protected]; Tel.: 86-10- Cuneo´ & Axsmith, which were previously placed in the 68999706; Fax: 86-10-68997803. “Taxodiaceae”, have been included.

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The Tiaojishan Formation (Fig. 2), near the village of Daohugou, is represented by volcaniclastic facies (Ren et al., 2002; Liu et al., 2004, 2010), deposited in a fluvio-lacustrine setting among the hills (Ren et al., 1996; Ren & Yin, 2003). The plant-fossiliferous bed (approximately 3 m) that lies in the lower part of the Formation consists of grey tuff (Ren et al., 2002). The Tiaojishan Formation lies between the Tuchengzi For- mation (Late Jurassic in age; above) and the Jiulong- shan Formation (Middle Jurassic in age; below) and is regarded to be Middle Jurassic to Late Jurassic (152– 168 Ma) in age based on fossil evidence of insects and conchostracans as well as radioisotopes (Shen et al., 2003; Chen et al., 2004; Liu et al., 2004, 2006, 2010; Wang et al., 2005; Zhou et al., 2007; Chang et al., Fig. 1. Map showing the fossil locality of Austrohamia acanthobractea 2009). sp. nov. in Daohugou of Inner Mongolia Autonomous Region, China. In the Daohugou flora, the other associated plant fossil remains includes Cladophlebis (Osmunda) Austrohamia is a monospecific genus (A. minuta sp., Anomozamites angulatus Heer, A. (Trymia) sp., Escapa, Cuneo´ & Axsmith) based on impressions rep- Cycadolepis sp., Ginkgoites sp., Pityospermus sp., resented by well-preserved leafy twigs and branches as Pityocladus sp., Zamites gigas (Lindley & Hutton) Mor- well as ovulate and pollen cones, from the Early Juras- ris, Yanliaoia sinensis Pan, Coniopteris burejensis (Za- sic of Chubut province, Argentina (Escapa et al., 2008a, lessky) Seward, Elatocladus sp., and Schizolepis dao- 2008b). hugouensis Zhang et al., which have been identified or In the present paper, a new species of Austrohamia described (Zhang, 2002; Ji et al., 2005; Zhang et al., from China, A. acanthobractea, sp. nov. from 10 spec- 2006, 2011; Zhou et al., 2007). imens, is described with discussion on its phytogeographical implications. 2 Results

1 Material and methods 2.1 Systematics Division Coniferophyta The specimens for the present investigation, in- Class Coniferopsida cluding ovulate cones along with vegetative branches Order Coniferales and twigs, were collected from the fossiliferous bed Family Cupressaceae sensu lato (41◦18N, 119◦14E) assigned to the Tiaojishan Forma- Genus Austrohamia Escapa, Cuneo´ & Axsmith 2008 tion, near the village of Daohugou, Ningcheng, Inner Species Austrohamia acanthobractea, Zhang, Mongolia Autonomous Region, China (Fig. 1). D’Rozario, Wang, Li & Yao, sp. nov. The fossils are preserved as compression- Specific diagnosis Leaves straight, or slightly falcate, impression specimens in a matrix of grey tuff, in which 3.0–7.0 mm long, 1.0–1.2 mm wide, with fairly thick the taphocoenosis is mostly dominated by the conchos- and distinct single vein; the vein sunken on upper sur- tracans. The specimens were analysed using a Zeiss MC face and raised on the lower one, especially near the 80 DX stereoscopic microscope (Carl Zeiss, Jena, Ger- basal part of leaf; leaf attached to axis by their decurrent many) with a camera lucida attachment. Photographs base, free part of the leaf extends outward at angles of were obtained under reflected light using a Nikon digi- 20–45◦; ovulate cone leathery, 5.0–8.0 mm long, 4.0– tal camera (Nikon, Kanagawa, Japan). 6.0 mm broad, bract broad triangular or semicircular, Micrographs of the fossils were made using a 3–5 mm long, 2–4 mm wide, inserted at an acute angle Jeol JSM-5610 LV scanning electron microscope (Jeol, to the axis, with spine-like head of 1.5–2.5 mm long, Tokyo, Japan). 1.0–1.5 mm wide, which is about one-third or half of Fossil specimens (MSF-Pb 1432–1441) are de- the whole length of the bract. Two oval ovules occur at posited in the repository of the Laboratory of Stratig- the proximal end of the adaxial surface of the bract. raphy and Paleontology, Institute of Geology, Chinese Derivation of specific epithet The specific epithet Academy of Geological Sciences, Beijing, China. acanthobractea is proposed to indicate the spine-like

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Fig. 2. Stratigraphic column of the Tiaojishan Formation, showing the layer with Austrohamia fossils. apex (acantho-) of the bract (bractea) of the ovulate leaves) (Fig. 3: B). In most cases, leaves on penulti- cone. mate and ultimate branches are straight, attached to the Holotype MSF-Pb 1432 axis by their decurrent base (Fig. 3: B; Fig. 4: A, B; Paratype MSF-Pb 1433 Fig. 5: A–I; Fig. 6: A, B). The free part of these leaves Locality Near Daohugou Village, Ningcheng County, extends outward at angles of 20–45◦ (mostly 30–40◦; Inner Mongolia Autonomous Region, China (Fig. 1) Fig. 5: A–E, H, I). In some cases, the free parts of the Stratigraphy Tiaojishan Formation (Fig. 2) leaves are slightly falcate, curving outward from the Age Middle Jurassic to Late Jurassic middle part, forming to a little larger angle of approx- Additional materials MSF-Pb 1434–1441 imately 50◦ to the axis (Fig 5: F, G). Leaves seem not displayed in a single plane, some leaves on the two sides 2.2 Description of the branches, others placed over the branches. Leaves Vegetative organs Vegetative branches are of at least are highly imbricate, especially near the internodes of two orders (incomplete). Ultimate branches 0.8–1.2 mm the distal portions of branches, where the leaves are wide, probably oriented in one plane, alternately or sub- shortened (Fig. 3: A; Fig. 4: A, B; Fig. 5: A; Fig. 6: oppositely arranged on penultimate branches which are A, B). 0.9–1.8 mm wide (Fig. 3: A). Leaves are helically ar- Ovulate cones Megasporangiate cones are elliptical, ranged. Individual leaves are dorsiventrally ﬂattened, small, 5.0–8.0 mm long and 4.0–6.0 mm wide, present linear-oblong with entire margins and rounded apexes singly or in clusters of two or three with each cone (Fig. 3: B; Fig. 4: A, B), 3.0–7.0 mm (mostly 4.0– borne at the end of a terminal branch (Fig. 3: A; Fig. 5: 6.0 mm) long and 1.0–1.2 mm (mostly 1.1 mm) broad. A, E). Sometimes they are found shed from branches Leaves with fairly thick and distinct single vein (midrib) (Fig. 3: A, C). When they occur in pairs, either both are which is slightly sunken on upper surface and raised terminally borne on the ultimate branches (Fig. 3: A; on the lower one (especially near the basal part of the Fig. 5: A), or the main axis ramiﬁes 5–10 mm below the

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Fig. 3. Austrohamia acanthobractea sp. nov., morphology of leafy branches and ovulate cones. A, MSF-Pb 1432. Holotype. Showing the alternate to sub-opposite disposition of last order branches, terminal clusters of ovulate cones; cone seems in one, two, or three in each branches, and each cone borne at the end of a terminal branch (arrow 1), sometimes shed (arrow 2); leaves helically arranged (arrow 3); the associated fossil conchostracans inthe specimen slab (arrow 4). Bar = 10 mm. B, Enlargement of part of the leafy branch, indicated by arrow 3 as in (A). Leaf straight, dorsiventrally flattened, linear-oblong, entire margins and rounded apexes (arrow 5), with fairly thick and distinct single vein and decurrent base (arrow 6). Leaf rib (Rib). Bar = 5 mm. C, Enlargement of an ovulate cone, indicated by arrow 1 in (A). Cone elliptical, ovuliferous complexes helically arranged, bracts leathery, apex rhomboidal and spine-like, two oval ovules (Ov) occur on the proximal part of the adaxial bract (Br) surface. Arrow 7, a cone scar. Bar = 5 mm. apex and both the new lateral branch and the main axis basal portion; the middle portion is expanded, shows a bear a terminal megasporangiate cone (Fig. 5: E). distinctive keel (convex) on the abaxial surface, which Ovulate cones are found either preserved as imma- is correspondingly concave. Even when two oval ovules ture (closed) ones bearing the ovules within (Fig. 5: E), occur at the proximal end of the adaxial surface of the or mature with ovules (seeds) shed (Fig. 3: C; Fig. 4: bract are counted, the presence of a major number cannot C, D; Fig. 6: C, D). In the immature ovulate cones, the be discarded (Fig. 3: C; Fig. 4: C; Fig. 6: C). bracts are adpressed to the cone axis (Fig. 5: E); but in the mature cones, the bracts are open, inserted at an acute angle to the cone axis (Fig. 3: C; Fig. 4: C, D; 3 Discussions Fig. 6: C, D). Ovuliferous complexes are helically and densely 3.1 Comparisons arranged along the cone axis (Fig. 3: C; Fig. 4: C, D; The new specimens (reconstructed in Fig. 7: A, B) Fig. 6: C, D). The bracts are leathery, lignified, broad from China having the vegetative characters, such as triangular or semicircular, 3–5 mm long and 2–4 mm alternate or sub-opposite disposition of the secondary wide. The apex of the bract is rhomboidal and spine- branches, helical phyllotaxy of leaves, lanceolate, dor- like, 1.5–2.5 mm long, 1.0–1.5 mm wide. In most cases, siventrally flattened, and univeined leaves with decur- the spine of the bract is approximately one-third of the rent base, together with the reproductive features, in- entire length of the bract; in other cases, the spine may cluding foliate nature of ovulate cones, apex of the reach to half the length of the whole bract (Fig. 3: C; bracts rhomboidal and spine-like, and ovules borne on Fig. 4: C, D; Fig. 6: C, D). The bract head is oriented the adaxial surface of bracts, indicate assignment to the nearly parallel to the cone axis (Fig. 3: C; Fig. 4: C, basal clades of the Cupressaceae sensu lato (formerly D; Fig. 6: C, D). The bracts may have a short stalk-like Taxodiaceae).

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Fig. 4. Austrohamia acanthobractea sp. nov., morphology of ovulate cone and leaf. A, B, Straight leaf with entire margins, rounded apexes (arrows 1, 3), and decurrent base (arrows 2, 4). C, D, Ovulate cone and its counterpart, showing the helically arranged ovuliferous complexes. Br, bract; Ov, ovule. A, C, D, Bar = 5mm; B, Bar = 1mm.

Within Cupressaceae sensu lato, morphological apex (Harris, 1943, 1953; Florin, 1958; Zhou, 1987; comparison places the Chinese fossil close to the ex- Yao et al., 1998; Fu et al., 1999; Farjon, 2005; Escapa tant genera Cunninghamia R. Br. and Taiwania Hayata, et al., 2008a). and the fossil genera Elatides, Sewardiodendron, and In the genus Cunninghamia, the leaves are not im- Austrohamia. The three mentioned fossil genera, in ad- bricated, margins serrulate and narrowed at base; in the dition to Cunninghamia and Taiwania, conform to a genera Taiwania, Elatides, and Sewardiodendron, the clade in the phylogeny of Escapa et al., 2008a. The leaves are imbricated, entire, and broad-based. closest comparison is with Austrohamia minuta. Leaves in the genera Taiwania and Elatides are 3.1.1 Main differences among the extant genera keeled and appressed to the branches, and rhomboidal Taiwania and Cunninghamia, and the fossil gen- in section; in the genus Sewardiodendron, the leaves are era Elatides, Sewardiodendron,andAustrohamia not keeled or appressed, and dorsiventrally ﬂattened. (Table 1) In the genus Austrohamia, ovuliferous Morphologically, genus Elatides are most similar cones are smaller, less than 1 cm both in length and to Athrotaxis (Sze & Li, 1963). Seward (1919) con- width, and the leaves have rounded apex; in the genera sidered that the former has the larger size of the seed Taiwania, Cunninghamia, Elatides, and Sewardioden- cones, and broader and thicker bract-scale complexes dron, the ovuliferous cones are larger, more than 1cm, than the latter; Harris (1953) thought that the ﬁnely ta- both in length and width, and their leaves have pointed pering apexes in Elatides are missing in Athrotaxis.

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Fig. 5. Austrohamia acanthobractea sp. nov.,morphology of vegetative branches, and the locations of the ovulate cones. A, MSF-Pb 1433. Paratype. The counterpart of holotype, showing the disposition of last order branches (1, 2), terminal clusters of ovulate cones, and the associated fossil conchostracans in the specimen slab (arrow 3). B–I, MSF-Pb 1434–1441. Last order leafy twigs bearing leaves (4, 5) that are not displayed in a single plane, some leaves on the two sides of the branches, whereas others placed over the branches. E, Megasporangiate cones borne in pairs at the apex of branches (arrows 6, 7), the main axis ramiﬁes 5–10 mm below the apex and both the new lateral branch and the main axis bear a terminal megasporangiate cone. F, G, Free part of the leaves slightly falcate, curve outward from the middle part, leads to a little larger angle of approximately 50◦ to the axis. Bar = 10 mm.

3.1.2 Comparison of morphological features of Yanliaoia Pan, a fossil taxodiaceous conifer genus, Austrohamia with Yanliaoia Pan and Elatocladus with only one species, Y. sinensis from the Late Jurassic (Halle) Harris From the Late Mesozoic of China, of Haifanggou Formation, in Western Liaoning, China two genera, Yanliaoia and Elatocladus, are also mor- (Pan, 1977), was believed to be closely allied to the phologically similar to Austrohamia. genus Elatides. It possesses the features of linear leaves

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Fig. 6. Scanning electron micrographs of Austrohamia acanthobractea sp. nov., showing morphology of ovulate cone and leaf. A, B, Straight leaf with entire margins, rounded apexes (arrows 1, 3), and decurrent base (arrows 2, 4). C, D, Ovulate cone, showing the helically arranged ovuliferous complexes. Br, bract; Ov, ovule. Bar scale = 1 mm. with entire margins, rounded apexes and distinct midrib, leaves are helically arranged about the stem, slightly and small female cones with its scales spirally arranged, decurrent, with a single, distinct midvein extending the spiny at their apex. All of these features are similar to length of each leaf, the form genus cannot be compared that in the genus Austrohamia. But in Yanliaoia, the to the natural genus of Austrohamia. features of the segment of its ultimate branches, the 3.1.3 Comparison with A. minuta The new species larger of its megasporangiate cones of more than 1 cm, (Figs. 3–7), which bears the structures of well-preserved the longer leaves of approximately 5–9 mm, the obtuse leafy twigs and branches as well as ovulate cones, shares pointed or narrow rounded apexes of its leaves, and the some common characteristics with A. minuta, such as: longitudinally arranged seeds in three ranks on each leafy shoots with at least two orders of branching, ul- scale are different to that of Austrohamia. timate branchlets alternate or sub-opposite with heli- Elatocladus Halle, a form genus, is applied to leafy cally arranged leaves which are dorsiventrally ﬂattened twigs with cuticle and reproductive parts unknown in and univeined; leaf base decurrent, apex rounded; and the order Coniferales (Gee, 1989; Sun et al., 2001). ovulate cones elliptical, less than 10 mm long, borne Although the leafy shoots of Elatocladus are morpho- terminally on ultimate and penultimate branches, com- logically similar to those of Austrohamia in that their posed of helically arranged bracts with ovules disposed

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ters is one of the synapomorphies of this basal Cu- pressaceae clade. These common characteristics suggest that the new plant can be assigned to the genus Austrohamia. The new species, however, differs from A. minuta in the morphology of the vegetative organs as well as the ovulate cones. In the vegetative features, the leaf of the new species is straight, 3.0–7.0 mm long, 1.0– 1.2 mm wide, compared to the falcate, smaller, 6.0– 7.0 mm long, 0.6–0.9 mm wide, leaf of A. minuta. Leaf of the new species is fairly thick with a distinct midrib, in contrast to the thin and slightly visible single vein in A. minuta. In the new species, the leaf is attached to the axis by a narrowed decurrent base, the free part of the leaf extends outward at a smaller angle of 20– 45◦, whereas in A. minuta, the leaf is adpressed to the axis for approximately half of its length then curves outward at a larger angle of 40–80◦. The new species has leathery, ligniﬁed, broad triangular or semicircular bract (Fig. 8: A) whereas, in A. minuta, cone bracts are weakly ligniﬁed and oval in shape (Fig. 8: B). The spine-like bract head (approximately one-third or half Fig. 7. Reconstruction of Austrohamia acanthobractea sp. nov. A, De- of whole bract length) of the new species is much more tailed reconstruction of an ultimate branchlet, showing the highly im- prominent than the rhomboidal bract apex (1/4 to 1/5 bricated leaves that are helically arranged on the axis. Bar = 5 mm; B, Reconstruction of the branch system showing terminal ovuliferous cones. the length of the bract length) of A. minuta. Bar = 10 mm.

3.2 Phytogeographical signiﬁcance on their adaxial surface. In A. minuta, ovulate cones The discovery of A. acanthobractea sp. nov. from have been found in pairs; in the new species there are the Middle to Late Jurassic of China provides impor- clusters of seed cones, similar to the cones observed tant phytogeographical implications for the cosmopoli- in the pollen cones of A. minuta; the presence of clus- tan distribution of the basal groups in Cupressaceae s.l..

Table 1 Comparison of morphological features of Austrohamia with the extant Taiwania and Cunninghamia, and the fossil Elatides and Sewardio- dendron Taiwania Cunninghamia Austrohamia Elatides Sewardiodendron Ultimate branches Orientation Irregular branching In whorls Oriented in one plane Irregular branching Not clear Arrangement Irregular Sub-opposite or in Alternate or Irregular Alternate or whorls sub-opposite sub-opposite Leaf Arrangement Helical Helical Helical Helical Helical, opposite or whorled Base Decurrent, broad Decurrent, narrowed Decurrent, narrowed Decurrent, broad Decurrent, broad Appressed Yes No No Yes No Keeled Yes No No Yes No Dorsoventrality Rhomboidal in section Dorsiventrally Dorsiventrally Rhomboidal in section Dorsiventrally flattened flattened flattened Shape Rhombic-ovate to Linear-lanceolate Linear-oblong Linear to triangular Lanceolate linear-lanceolate Margin Entire Serrulate Entire Entire Entire Apex Obtuse or acute Acute Rounded Pointed Pointed Imbricate Yes No Yes Yes Yes Ovulate cone Shape Ovate to elongated Ovoid or rounded Elliptical Elliptical Ovate to elongated Length (cm) >1 >1 >0.5, <1 >1 >1 Reference Fu et al., 1999; Fu et al., 1999; Escapa et al., 2008a Harris, 1943, 1953; Florin, 1958; Yao Farjon, 2005 Farjon, 2005 Zhou, 1987 et al., 1998

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seed cones of Pararaucaria Wieland (Stockey, 1977) and Romeroites Spegazzini (1924) from Argentina, the woods of Prototaxodioxylon Vogellehner, Taxodioxy- lon Hartig, Protocupressinoxylon Eckhold, and Cu- pressinoxylon Goeppert from Laurasia (Zhou & Zhang, 1989; Philippe, 1994), compressed/impressed reproductive and/or vegetative remains of Farndalea Bose, Se- quoia Endl., Sewardiodendron Florin, the form genus of Cunninghamites Presl, Sciadopitytes Goeppert and Menge, and Elatocladus Halle. All of these genera have Fig. 8. Comparison of the morphology of the ovuliferous complexes a possible taxodiaceous affinity (Florin, 1922, 1958; (schematic drawings). A, Austrohamia acanthobractea sp. nov.; B, A. Harris, 1943, 1979; Endo, 1951; Bose, 1961). But most minuta (redraw from Escapa et al., 2008a). Br, bract. Bar = 2 mm. of these members referred to Taxodiaceae are based only on fragmentary remains (Yao et al., 1998). Based on the fossil records, the basal groups in Cu- By the Cretaceous, the record of fossil Cupres- pressaceae s.l. (previously Taxodiaceae) originated dur- saceae had reached worldwide distribution, suggesting ing the Triassic and were well established by the Jurassic that certain features were already present before that (Yao et al., 1998). Although no clear evidence suggests time (Taylor et al., 2009). that the extinct family Voltziaceae from Late Permian The new discovery of A. acanthobractea sp. nov., and Triassic is a natural group, this family was thought from the Middle to Late Jurassic of China, is a new to be the ancestor of the Cupressaceae s.l. (Miller, 1977, record in the Northern Hemisphere (on the opposite 1982; Stockey et al., 2005). This family was widespread side of the Pacific). in both the Southern Hemisphere, such as conifer seed All of these descriptions would suggest the cos- cones of Voltziopsis Potonie,´ Rissikia Townrow, and mopolitan distribution of the basal groups in Cupres- Telemachus Anderson from the Triassic of Gondwana saceae, from the Early Jurassic to Late Jurassic, before (Townrow, 1967a, 1967b, 1969; Yao et al., 1997; Es- the disaggregation of the Pangaea. Exchanges between capa et al., 2010), and the Northern Hemisphere, such as East Asia and South America were possible by way of the permineralized seed cone of Pseudovoltzia liebeana the land-bridge. The basal groups in the family Cu- (Geinitz) Florin from the Permian of Rhineland, Ger- pressaceae s.l. (formerly Taxodiaceae), would migrate many (Schweitzer, 1963; Yao et al., 1997). The oldest, from southern America, and through northern Africa, to generally accepted fossil assignable to the Cupressaceae Europe, and then to East Asia. s.l.,isParasciadopitys aequata Yao, Taylor & Taylor, a taxodiaceous seed cone, from the early Middle Triassic Acknowledgements We express our gratitude to of Fremouw Peak, Antarctica (Yao et al., 1993, 1997; Professor Cheng-Sen LI, Institute of Botany, Chinese Taylor et al., 2009; Escapa et al., 2010). Academy of Sciences, for giving us advice about From South America, A. minuta from the Early the identification of the specimens. We thank Sheng- Jurassic of Chubut province, Argentina, which has ovu- Hui DENG, Laboratory Center, Research Institute of late and pollen cones along with vegetative branches Petroleum Exploration and Development, for providing and twigs preserved (Escapa et al., 2008a, 2008b), was relevant references. This research was supported by the the first record of Cupressaceae s.l. that shows affinity China Geological Survey (Grant Nos. 1212010911047 with the extant genera Taiwania and Cunninghamia in and 1212010811057). The authors acknowledge the basal Cupressaceae (formerly Taxodiaceae). support from the Ministry of Science and Technology, Elatides and Sewardiodendron, the only two tax- China (Grant No. 2006FY120300–12). odiaceous genera from the Middle Jurassic, also have ovulate and pollen cones as well as vegetative twigs preserved. Elatides williamsoni Lindley & Hutton and References the vegetative remains of Sewardiodendron laxum were found from Yorkshire, UK (Harris, 1943, 1979; Escapa Bose MN. 1961. Leaf-cuticle and other plant microfossils from et al., 2008a); compressed seed and pollen cones along the Mesozoic rocks of Andøya, Norway. Paleobotanist 8: 1–7. with leafy shoots of S. laxum were discovered from Brunsfeld SJ, Soltis PS, Soltis DE, Gadek PA, Quinn CJ, Strenge Yima, Henan, central China (Yao et al., 1998). DD, Ranker TA. 1994. Phylogenetic relationships among In the Jurassic, other genera from both the Southern the genera of Taxodiaceae and Cupressaceae: Evidence from and Northern Hemispheres include the permineralized rbcL sequences. Systematic Botany 19: 253–262.

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