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A New Species of Aneurophyton (Progymnospermopsida) from the Middle Devonian of West Junggar, Xinjiang, China, and Its Paleophytogeographical Significance

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A New Species of Aneurophyton (Progymnospermopsida) from the Middle Devonian of West Junggar, Xinjiang, China, and Its Paleophytogeographical Significance Int. J. Plant Sci. 174(8):1182–1200. 2013. ᭧ 2013 by The University of Chicago. All rights reserved. 1058-5893/2013/17408-0007$15.00 DOI: 10.1086/671807 A NEW SPECIES OF ANEUROPHYTON (PROGYMNOSPERMOPSIDA) FROM THE MIDDLE DEVONIAN OF WEST JUNGGAR, XINJIANG, CHINA, AND ITS PALEOPHYTOGEOGRAPHICAL SIGNIFICANCE Qing Jiang,*,† Yi Wang,* Hong-He Xu,1,* and Jing Feng† *State Key Laboratory of Paleobiology and Stratigraphy, Nanjing Institute of Geology and Paleontology, Chinese Academy of Sciences, 210008 Nanjing, People’s Republic of China; and †University of Chinese Academy of Sciences, 100049 Beijing, People’s Republic of China Aneurophyton doui sp. nov. (Aneurophytales, Progymnospermopsida) is described from the late Middle Devonian Hujiersite Formation of Hoxtolgay, Xinjiang, northwest China, as a plant with at least four orders of axes and ultimate units (vegetative appendages/fertile organs). Spines cover the surface of all orders of axes and ultimate units. The second-order axes and the vegetative appendages are closely inserted in pairs and helically arranged. From the second order, axes of subsequent orders are produced by bifurcation. The veg- etative appendages are unwebbed and up to three times dichotomous. The fertile organ is pinnate, and the fertile organ axes are opposite or subopposite with adaxial, elliptical sporangia. This is the first formal report of Aneurophyton beyond the coasts of the Rheic Ocean. West Junggar, the locality of present species of Aneurophyton, played a key role in the dispersal of Aneurophytales in the Middle Devonian. Keywords: Aneurophyton, progymnosperm, Devonian, paleophytogeography, Xinjiang. Introduction of Aneurophyton, and the paleophytogeographical significance and possible migration route of Aneurophytales are discussed. The Progymnospermopsida, being characterized by pterid- ophytic free-sporing reproduction but gymnospermous sec- ondary vascular tissues (Beck 1960; Bonamo 1975; Gensel and Material and Methods Andrews 1984; Hammond and Berry 2005), is subdivided into three orders: Aneurophytales, Archaeopteridales, and Protop- Twenty-three specimens were collected from a small quarry ityales (Kra¨usel and Weyland 1941; Beck 1960; Beck and by National Highway G217, ∼20 km north of the town of Wight 1988). Among them, Aneurophytales is thought to be Hoxtolgay, Hobuksar Menggol Autonomous County, Xin- the most primitive (Kra¨usel and Weyland 1941; Beck 1960; jiang, China (lat. 46Њ3655N, long. 86Њ0105E). The strata Hammond and Berry 2005). Genera in Aneurophytales estab- belong to the G217 Highway Section of the Upper Member lished on the basis of both morphological and anatomical char- of the Hujiersite Formation (for detailed introduction and fig- acters include Tetraxylopteris Beck 1957, Rellimia (Dawson) ures, see Xu et al. 2012a, forthcoming). Leclercq et Bonamo 1973, and Aneurophyton Kra¨usel et Wey- Specimens are preserved as coalified compressions and im- land 1923. Triloboxylon Matten et Banks 1966 is mainly based pressions in yellow-green to gray tuffaceous siltstone. The on anatomy but also some morphological characters. Other plant fossil bed was dated to the late Mid-Devonian (Givetian) genera, such as Proteokalon Scheckler et Banks 1971, Rei- on the basis of palynology (Xu et al., forthcoming). Plants mannia Arnold 1935, and Cairoa Matten 1973, were estab- reported from the Hujiersite Formation include Lepidoden- lished on anatomical characters only (Scheckler 1975; Beck dropsis theodori Jongmans (Cai and Wang 1995), Tsaia conica and Wight 1988; Hammond and Berry 2005). Aneurophyta- Wang et al. (2004), Leclercqia cf. complexa Banks et al. (Xu lean rhizomes were recently described from Gilboa, although and Wang 2008), Leclercqia uncinata Xu et al. (2011), Has- no genus was designated (Stein et al. 2012). kinsia hastata Berry et Edwards, Haskinsia sagittata Edwards The diagnostic characters of Aneurophyton come mainly et Benedetto (Xu et al. 2008), Compsocradus givetianus from the type species, A. germanicum Kra¨usel et Weyland (Wang) Fu et al. (2011), and Hoxtolgaya robusta Xu et al. 1923, widely reported from the Middle Devonian of Eur- (2012b). america (Serlin and Banks 1978; Schweitzer and Matten 1982). Sharp needles were used to remove the matrix embedding In this article, A. doui sp. nov. is described from the late Middle the plant fossils to obtain morphological information (de´gage- Devonian Hujiersite Formation of West Junggar, Xinjiang, ment; Fairon-Demaret et al. 1999). Macrophotographs were China. This locality is far away from the earlier occurrences taken using a Nikon D-100 camera with a Nikkor 105-mm macro lens; cross polarized illumination was used to enhance 1 Author for correspondence; e-mail: [email protected]. contrast. Microphotographs were taken using a Leica MZ-16 Manuscript received October 2012; revised manuscript received April 2013; stereomicroscope with annular ring illumination and a Leica electronically published September 4, 2013. D480 digital camera. 1182 This content downloaded from 119.78.212.207 on Sat, 21 Dec 2013 04:16:45 AM All use subject to JSTOR Terms and Conditions Fig. 1 Aneurophyton doui sp. nov. from the Middle Devonian Hujiersite Formation, Xinjiang, China. A, Holotype and paratype occur in one rock. Vegetative (left, holotype, PB21568) and fertile part (right, paratype, PB21569A) of the plant. Scale bar p 10 mm. B, Enlargement of the portion indicated by arrowhead 2 in A, showing four second-order axes inserted into the first-order axis. Note that the lower pair (IIA1 and IIA2) extends opposite the observer; the upper pair (IIB1 and IIB2) extends toward the observer. IIA1 dichotomizes into two third-order axes p (III1 and III2). Scale bar 10 mm. C, Enlargement of the portion indicated by arrowhead 3 in A, showing bifurcation of a second-order axis. Spines are on both second-order and third-order axes (arrowheads). Scale bar p 5 mm. D, Enlargement of the portion indicated by arrowhead 1inA, showing spines (arrowheads) and longitudinal coal-filled depression on the surface of the axis. Scale bar p 5 mm. E, F,Partand counterpart. Two isolated terminal axes with vegetative appendages and detached sporangia at the distal part. Scale bar p 10 mm. PB21570. G, Enlargement of the portion indicated by the black arrowhead in F, showing tongue-shaped depressions (arrowheads) at the basal part of vegetative appendages indicating embedded appendages in one pair. Scale bar p 2 mm. H, Arrowed axis in F, after preparation. Note that the paired vegetative appendages are helically attached. Scale bar p 10 mm. This content downloaded from 119.78.212.207 on Sat, 21 Dec 2013 04:16:45 AM All use subject to JSTOR Terms and Conditions Fig. 2 Aneurophyton doui sp. nov. from the Middle Devonian Hujiersite Formation, Xinjiang, China. A, B, Part and counterpart. Note the four orders of axes (I–IV) and vegetative appendages attached to the fourth-order axes. Scale bar p 10 mm. PB21571. C, First-order axis in B, before preparation. The tongue-shaped depression (black arrowhead) indicates the attachment of another second-order axis. Scale bar p 10 mm. D, Enlargement of the third- and fourth-order axes in A, after preparation. Paired vegetative appendages are attached helically to the fourth-order axes. Scale bar p 10 mm. E, Enlargement of a pair of vegetative appendages (black arrowhead in D), showing the paired appendages dichotomizing three times into the flattened segments. The first dichotomy of the complete appendage results in the left branchlet (black arrowhead) and the right branchlet (white arrowhead). Note that the left branchlet leans into matrix, the right parallel to the slab surface. Scale bar p 5 mm. F, Enlargement of a portion of the fourth-order axis IV4 (white arrowhead in A), showing the decurrent bases of vegetative appendages and the spines (white arrowheads) on the axis. Scale bar p 5 mm. G, Enlargement of the distal portion of axis III in A, showing the third-order axis dichotomizing into two fourth-order axes. Spines are on both orders (white arrowheads). Tongue-shaped depressions are seen at the basal part of the vegetative appendage p (black arrowhead). Scale bar 5 mm. H, Enlargement of a portion of axis IV4 (black arrowhead in A). Spines are at the base of the sterile unit (white arrowheads). Tongue-shaped depressions indicate the embedded vegetative appendages (black arrowheads). Scale bar p 2mm. This content downloaded from 119.78.212.207 on Sat, 21 Dec 2013 04:16:45 AM All use subject to JSTOR Terms and Conditions JIANG ET AL.—MID-DEVONIAN ANEUROPHYTON FROM CHINA 1185 Systematic Paleobotany Class—Progymnospermopsida Beck 1960 Order—Aneurophytales Kra¨ usel et Weyland 1941 emend. Beck 1960 Family—Aneurophytaceae Kra¨usel et Weyland 1941 Genus—Aneurophyton Kra¨usel et Weyland 1923 emend. Schweitzer et Matten 1982 Type Species—Aneurophyton germanicum Kra¨usel et Weyland 1923 emend. Schweitzer et Matten 1982 Species—Aneurophyton doui Jiang, Wang, Xu et Feng sp. nov. Diagnosis. At least four orders of axes and ultimate units (vegetative appendages/fertile organs) are known. Spines, 0.5– 1.5 mm long, occur sparsely on the surface of all orders of axes and ultimate units. First-order axes, up to 6 mm wide and at least 90 mm long, demonstrate only a slight distal taper. Second-order axes, up to 4 mm wide, diverge helically from the first-order axis in closely inserted pairs. From the second order, axes of subsequent orders are produced by bifurcation; third-order axes are 2.0–2.5 mm wide, and fourth-order axes are 1.5–2.0 mm wide. Vegetative appendages, 7.0–13.0 mm long as a whole, diverge from the third- or fourth-order axes in helical pairs, as a one to three times dichotomizing system of three-dimensionally-extended flattened branchlets. Fertile organs, 3.7–8.0 mm long as a whole and borne on the third- or fourth-order axes, are up to three times opposite/sub- opposite pinnate systems. All three orders of fertile organ axes, in most cases recurved and rarely straight, bear sporangia or organ axes.
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