1. Introduction
Middle Devonian herbaceous lycopsids are widely accepted to have palaeobiogeographical significance if their leaf morphology has been correctly worked out and documented allowing accurate generic and specific determination (Bonamo, Banks & Grierson, Reference Bonamo, Banks and Grierson1988). There remain a few cases where historically important lycopsid discoveries have yet to be reinvestigated to determine their exact leaf morphology. One example is the report of Protolepidodendron scharyanum Krejči (the name should be P. scharianum Kräusel et Weyland, see Xu & Wang, in press) from Xinjiang, NE China. Specimens originally determined by Sze (Reference Sze1960, Reference Sze1961) on the basis of once-divided leaves were later re-assigned to P. cf. cathaysiense Schweitzer et Cai by Cai & Wang (Reference Cai, Wang and Li1995) who noticed one leaf that appeared to trifurcate. The question of the exact determination was left ambiguous as no spines were visible on the margin of leaves.
The identification of these specimens is important because of their palaeogeographical situation. Protolepidodendron Krejči ex Gothan (for authorship see Xu & Wang, in press) is generally held to refer to a herbaceous lycopsid with once-bifurcate leaves (Gothan, Reference Gothan1921, pp. 225–6). Most specimens previously attributed to Protolepidodendron have been reassigned to Leclercqia Banks, Bonamo et Grierson (Reference Banks, Bonamo and Grierson1972), which has a five-tipped leaf with a prominent central elongate segment and two pairs of shorter lateral segments (Banks, Bonamo & Grierson, Reference Banks, Bonamo and Grierson1972; Fairon-Demaret, Reference Fairon-Demaret1974, Reference Fairon-Demaret1980, Reference Fairon-Demaret1981), and is found widely in Laurentia and Gondwana (Meyer-Berthaud et al. Reference Meyer-Berthaud, Fairon-Demaret, Steemans, Talent and Gerrienne2003). However Protolepidodendron from Yunnan, South China (Halle, Reference Halle1936; Schweitzer & Cai, Reference Schweitzer and Cai1987) was transferred to Minarodendron Li (Reference Li1990), characterized by trifurcate spiny leaves. Minarodendron (Protolepidodendron) cathaysiense Li (Reference Li1990) is restricted to South China, the only plate yet to show a strongly endemic Middle Devonian macroflora (Wang et al. Reference Wang, Berry, Hao, Xu and Fu2007). The palaeogeographical positions of the Kazakhstania plates, on one of which the locality was situated (Liao & Ruan, Reference Liao, Ruan, Zhou and Lin1995), in the Middle Devonian is problematical (Scotese & McKerrow, Reference Scotese and McKerrow1990; Cocks & Torsvik, Reference Cocks and Torsvik2002; Torsvik & Cocks, Reference Torsvik and Cocks2004). This investigation is to demonstrate the true leaf morphology of Sze's (Reference Sze1960, Reference Sze1961) specimens and therefore to determine if they show affinity to the more or less cosmopolitan herbaceous lycopsid flora or the endemic flora of Yunnan.
2. Material and methods
The detailed locality and horizon was not stated by Sze (Reference Sze1960, Reference Sze1961), according to whom all materials were collected from the Hoxtolgay area, Hoboksar Mongol Autonomous County, North Xinjiang, NW China. The fossil-bearing horizon also yielded Hostimella sp., Lepidodendropsis? (Colpodexylon?) arborescens Sze, and Lepidodendropsis? dzungariensis Sze. The horizon is inferred to be the Hujiersite Formation (Givetian) from this fossil plant assemblage (Cai & Wang, Reference Cai, Wang and Li1995). A few localities (Fig. 1) are probably involved. More recently reported plants from the Hujiersite Formation of the Hoxtolgay area include: Sawdonia curstipa Wang & Hao (Reference Wang and Hao1996), Haskinsia sagittata (Cai & Wang, Reference Cai, Wang and Li1995; Xu et al. in press), H. hastata (Xu et al. in press), Colpodexylon gracilentum (H.-H. Xu, unpub. Ph.D. thesis, Nanjing Institute of Geology and Palaeontology, 2006), Tsaia conica (Wang et al. Reference Wang, Xu, Fu and Tang2004), and Compsocradus sp. (H.-H. Xu, unpub. Ph.D. thesis, Nanjing Institute of Geology and Palaeontology, 2006).
![](https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20180417041851260-0410:S0016756807004256:S0016756807004256_fig1g.gif?pub-status=live)
Figure 1. Generalized geological and geographical map showing probable localities of Sze's (Reference Sze1960, Reference Sze1961) collection in Xinjiang, NW China, and the relative generalized position of the Yunnan Middle Devonian flora.
A total of six specimens (PB3246–PB3251) attributed to Protolepidodendron scharyanum by Sze are preserved as compressions and deposited at Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences. A short leafy stem on the reverse side of Sze's specimen PB3246 was uncovered using serial dégagement (Fairon-Demaret, Hilton & Berry, Reference Fairon-Demaret, Hilton, Berry, Jones and Rowe1999). Leaves on the counterpart to Sze's specimen PB3246 (PB3247) are also illustrated. Macrophotography was carried out using a Nikon D100 digital camera with cross-polarized illumination.
3. Description
The new fragmentary unbranched axis with persistent leaves described here is 22mm long and 3.8–4.2mm wide (Fig. 2a). The surface of the stem was highly compressed, so that it is hard to recognize patterns of leaf bases. Leaves in profile are linear with a swollen base, showing an abaxially recurved tip (Fig. 2a, leaves 1, 3 and 4) distal to a division, or a division to form two or more segments (Fig. 3). The angle between leaf and stem ranges from 40° (Fig. 2a) to 50° (Fig. 2b).
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Figure 2. Leclercqia cf. L. complexa from the Middle Devonian of North Xinjiang, China. (a) New discovered short leafy stem on the reverse side of PB3246 (Sze, Reference Sze1960, Reference Sze1961). (b) Enlargement of leaves in Sze's unprepared specimen PB3247, upper leaf partially prepared, four segments visible distally (arrow); lower unprepared leaf shows typical bifurcation as observed in Protolepidodendron scharyanum Kräusel et Weyland. (c, d) Stages of serial dégagement to the leaf 2 in (a). Lettering of segments follows Fairon-Demaret (Reference Fairon-Demaret1980). All scale bars = 1mm.
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Figure 3. Leclercqia cf. L. complexa from the Middle Devonian of North Xinjiang, China. Line drawings of leaves in Figure 2. (a) Arrowed leaf in Figure 2b; (b,c,d) correspond to leaves 5, 4 and 3 in Figure 2a; (e,f,g) three stages of serial dégagement to the leaf 2 in Figure 2a (f and g correspond to Fig. 2c and d). Lettering of segments follows Fairon-Demaret (Reference Fairon-Demaret1980). Scale bar = 2mm.
Serial dégagement was performed on leaf 2 (Fig. 2a). Like most partly prepared leaves, this leaf shows three segments (b, c and d in Fig. 3e) before dégagement. In addition, an obvious wider segment was buried in the matrix under the segment d (Fig. 3e). After removing segment d, the long central segment E was finally exposed (Fig. 2c; Fig. 3f). Segment a was also revealed after segment b was removed (Fig. 2d; Fig. 3g).
The unforked part of the leaf is 0.33–0.41mm wide. The leaf divides 3.4–3.8 (mean 3.61, n=4)mm from the base, forming a long central segment (up to 3.1mm long, Fig. 2a, leaf 3; Fig. 3d) recurved abaxially and a pair of lateral laminae that each bifurcates proximally into two slender segments 0.77–1.54mm in length. The arrangement of all five segments is always three-dimensional. It is impossible to discern the proximal ends of all segments. In some leaves, four distal segments could be seen at the same time, e.g. Sze's (Reference Sze1960, Reference Sze1961) specimen PB3247 (Fig. 2b, arrow; Fig. 3a); leaf 5 on the new stem (Fig. 2a; Fig. 3b).
4. Discussion
Bifurcate leaves were for a long time accepted as a distinctive character of the genus Protolepidodendron. Schweitzer & Cai (Reference Schweitzer and Cai1987), for the first time, noted the trifurcate and spiny microphylls in specimens previously attributed to P. scharyanum by Halle (Reference Halle1936) from the Middle Devonian of Yunnan, South China, and placed the specimens in a new species, P. cathaysiense, effectively enlarging the generic concept. Additional specimens of P. cathaysiense were studied by Li (Reference Li1990). Multiperforate bordered pits in the tracheids, as well as the trifurcate leaves, supported transfer to the new genus Minarodendron Li. Minarodendron is widely accepted as an endemic lycopsid that is probably related to Leclercqia (Kenrick & Crane, Reference Kenrick and Crane1997). Therefore Protolepidodendron is no longer recognized in South China.
Cai & Wang (Reference Cai, Wang and Li1995) re-examined Xinjiang specimens attributed to P. scharyanum, from which three leaf distal segments were recognized (Cai & Wang, Reference Cai, Wang and Li1995, pl. 16, figs 1, 2). Specimens were transferred to Protolepidodendron cf. P. cathaysiense. However we recognize three-dimensional, five-tipped leaves from the newly studied leafy stem, and the specimens should therefore be reassigned to Leclercqia.
The type species, L. complexa, at the type locality in New York State, was measured as: stem 3.5–7.0mm wide, leaf 4.0–6.5mm long as a whole and bearing one-half to two-thirds of its length a pair of lateral divisions. The central and the lateral segments of the leaf are 2.5–3.0mm and 1.0–2.0mm long respectively (Banks, Bonamo & Grierson, Reference Banks, Bonamo and Grierson1972). These measurements are well matched with dimensions of the present plant. No information about the ligule, sporophylls, sporangia, spores or anatomy was available, so, it is proper to attribute the present plant to Leclercqia cf. L. complexa (cf. Berry, Reference Berry1994). L. complexa differs from a second species, L. andrewsii Gensel et Kasper, from the Lower Devonian of New Brunswick, Canada (Gensel & Kasper, Reference Gensel and Kasper2005), in that the median and lateral segments of the leaf in the latter are all in one plane and adaxially recurved.
In view of the above, Sze's (Reference Sze1960, Reference Sze1961) specimens of P. scharyanum, assigned to P. cf. cathaysiense by Cai & Wang (Reference Cai, Wang and Li1995), are hereby transferred to L. cf. complexa. This is the first confirmed record of Leclercqia on the basis of demonstration of leaf morphology in Chinese Middle Devonian strata.
Until now, the Middle Devonian lycopsid flora of North Xinjiang contains Haskinsia sagittata Edwards et Benedetto (Cai & Wang, Reference Cai, Wang and Li1995), H. hastata Berry et Edwards (Xu et al. in press), Colpodexylon Banks, Archaeosigillaria Kidston (Dou et al. Reference Dou, Sun, Wu and Gu1983) and Leclercqia (this work). The floral assemblage shows much more similarity to that of North America (see Grierson & Banks, Reference Grierson and Banks1963, Reference Grierson and Banks1983) and Venezuela (see Berry, Casas & Moody, Reference Berry, Casas and Moody1993; Berry & Edwards, Reference Berry and Edwards1995, Reference Berry and Edwards1996) than to that of the Xichong Formation, Yunnan (Wang et al. Reference Wang, Berry, Hao, Xu and Fu2007). The only exception is an unusual form of Colpodexylon in Yunnan (Schweitzer & Cai, Reference Schweitzer and Cai1987), new specimens of which show no Colpodexylon (‘embayed xylem’ – Banks, Reference Banks1944) structure and which therefore may be transferred to a new genus (Y. Wang et al. unpub. data). The transfer of P. cf. cathaysiense to Leclercqia therefore emphasizes the palaeophytogeographical separation of North Xinjiang from Yunnan.
Since its establishment, Leclercqia has been reported from the Early to Middle Devonian more or less worldwide, including the type locality in New York State, USA (Banks, Bonamo & Grierson, Reference Banks, Bonamo and Grierson1972), Australia (Fairon-Demaret, Reference Fairon-Demaret1974; Meyer-Berthaud et al. Reference Meyer-Berthaud, Fairon-Demaret, Steemans, Talent and Gerrienne2003), Germany (Fairon-Demaret, Reference Fairon-Demaret1980), Belgium (Fairon-Demaret, Reference Fairon-Demaret1981), Venezuela (Berry, Reference Berry1994), New Brunswick, Canada (Gensel & Kasper, Reference Gensel and Kasper2005; Gensel & Albright, Reference Gensel and Albright2006) and Xinjiang, China (this work). The position and boundaries of the plates that later make up the Kazakhstania plate are uncertain in Middle Devonian time (Torsvik & Cocks, Reference Torsvik and Cocks2004). Middle Devonian plant localities in North Xinjiang and Kazakhstan were potentially close or even on the same microplate (Liao & Ruan, Reference Liao, Ruan, Zhou and Lin1995) and have similar members of Middle Devonian flora (Ananiev, Reference Ananiev1960; Jurina, Reference Jurina1988; Cai & Wang, Reference Cai, Wang and Li1995). It is likely that some Kazakhstan specimens currently attributed to Protolepidodendron may be transferred to Leclercqia after further investigation.
Evidence from spores supports the occurrence of Leclercqia in North China. As the dispersed spore Dibolisporites heitaiensis Ouyang (Reference Ouyang1984) from the Middle Devonian of Heilongjiang, NE China, shows great similarity to the in situ spore of L. complexa (Richardson, Bonamo & McGregor, Reference Richardson, Bonamo and McGregor1993), Ouyang (Reference Ouyang1984) suggested that megafossils of Leclercqia would probably be discovered from the Middle Devonian of North China as well as Siberia. Geng (Reference Geng1992) reported L. complexa from the Lower Devonian (Pragian) of Sichuan, South China. But the assignment was rejected (Cai & Wang, Reference Cai, Wang and Li1995) because of the lack of typical features of Leclercqia.
When marked on the Middle Devonian paleogeographical and palaeoclimatic reconstruction, occurrences of Leclercqia are widely distributed, most of which are in the arid climatic belt (Meyer-Berthaud et al. Reference Meyer-Berthaud, Fairon-Demaret, Steemans, Talent and Gerrienne2003). Leclercqia in Xinjiang suggests a tropical distribution. Considering the occurrence also in the warm temperate climatic belt (Venezuela: Berry, Reference Berry1994), we suggest that Leclercqia is more or less cosmopolitan in the Middle Devonian, with the only notable exception being South China, rather than distributed in the same so-called arid climatic belt as proposed by Meyer-Berthaud et al. (Reference Meyer-Berthaud, Fairon-Demaret, Steemans, Talent and Gerrienne2003).
Acknowledgements
We thank Dr Wang Q., Institute of Botany, Chinese Academy of Sciences, for helpful suggestions. The work is supported by National 973 Project (No. 2006CB806400) and NSFC No. J0630967.