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Reproductive Biology of the Arborescent Seed-Fern Linopteris Obliqua

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Reproductive Biology of the Arborescent Seed-Fern Linopteris Obliqua DOI: 10.2478/fbgp-2020-0002 E. L. Zodrow, J. Pšenička & Wei-Ming Zhou, Reproductive biology of the arborescent seed-fern Linopteris obliqua REPRODUCTIVE BIOLOGY OF THE ARBORESCENT SEED-FERN LINOPTERIS OBLIQUA: IMPLICATIONS FOR TAXONOMY (MEDULLOSALES, LATE PENNSYLVANIAN SYDNEY COALFIELD, CANADA) Erwin L. Zodrowa, Josef Pšeničkab & Wei-Ming Zhouc a 503 Coxheath Road, Sydney, Nova Scotia, Canada; e-mail address: [email protected] b Centre of Palaeobiodiversity, West Bohemian Museum in Pilsen, Kopeckého sady 2, Pilsen 30100, Czech Republic, [email protected] c State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology and Center for Excellence in Life and Paleoenvironment, Chinese Academy of Sciences, Nanjing 210008, China; e-mail address: [email protected] Abstract: A shaley slab (65 x 45 x 7 cm) from the Sydney Coalfield, Canada, Cantabrian age, on splitting apart revealed 2 – 3 layers each entombing thousands of abscised pinnules of Linopteris obliqua and eight dispersed compound-synangial structures. The campanulary-ventral-sporal micromorphology of the best preserved structure of these compares sufficiently well with previously reported structures from the Sydney Coalfield named Potoniea krisiae. Earlier studies involving larger sampling suites furthermore contributed to the observation that Hexagonocarpus sp. (female organ) and P. krisiae (male organ) usually co-occur with abscised L. obliqua pinnules; however, these two organs do not co-occur on isochronous bedding planes. In the absence of confirmatory organic attachments, the presented data provide as yet the strongest support for the hypothesis of the organs’ connectivity, but whether female-male trees existed or not, and the mode of attachment of the organs remain unknown. Hypothesized for the latter is pinnate attachment. Keywords: Medullosalean, Potoniea krisiae, Hexagonocarpus INTRODUCTION The species’ concept of Linopteris (ex Dictyopteris) extensive coal diamond-drill programs, commented obliqua (Bunbury 1847) is based on a single pin- on the caducous nature of the pinnules, and estab- nule specimen, lectotype “186” (so designated by lished, amongst others, the L. obliqua Zone. It is Zodrow et al. 2007). It is deposited with Cambridge emended through the collection of L. obliqua spec- University, Botany School, together with Bun- imens from the uppermost strata at Point Aconi, bury’s hand-written “Catalogue of Fossil Plants in which is part of Bell’s originally defined Ptychocar- my Collection, 1850; Part 1, Ferns and Lycopodi- pus unitus Zone (Zodrow & McCandlish 1978). aceae” in which he recorded two additional pin- Bell’s Zonation is used by Zodrow & Cleal (1985, nules of this species “187 and 188” from Cape Bre- fig. 5, among other plant fossils) for correlating ton, Nova Scotia (cf. Zodrow et al. 2007). In fact, non-marine Asturian (Westphalian D) and Can- Bunbury unknowingly brought to the fore the tabrian strata from Europe and North America. autecological nature of this species. All three pin- In general, L. obliqua is restricted to tropical nules originated from what is now called the Har- Euromerica of Late Pennsylvanian age as is sum- bour Seam of the Sydney Mines Formation of the marized by Tenchov & Popov (1987) and Zodrow Late Pennsylvanian Sydney Coalfield, Canada & McCandlish (1978), but it also occurs in the (Figure 1C). Mazon-Creek Flora, U.S.A. (Pfefferkorn 1979). However, Bell (1938, fig. 1, pt.1) mapped the The present contribution focuses on, and is com- stratigraphic occurrences and the lateral extent of plementary to earlier studies to strongly support L. obliqua in the Sydney Coalfield based on the the connectivity of the male organ Potoniea krisiae 13 Folia vol. 54, No. 1–2, 2020 Pšenička et al. 2019 and the female organ Hexagono- MATERIAL, METHODS, carpus sp., compared with H. crassus Renault AND STRATIGRAPHIC AGE 1890 by Zodrow et al. (2007), to the seed-fern plant L. obliqua (Zodrow 2002, 2007). Thus, Bell’s origi- The study material, accession number 08-10/15-9, nal L. Zone is likely characterized by a natural originated from a 65 x 45 x 7 cm in situ argilla- species, not known for any other Carboniferous- ceous-shaley slab 1-2 m above the Hub Seam in zone phytostratigraphy. In effect, illustrated is the the now-closed (2002) submarine Prince Mine near utility of large specimens collected with time- Point Aconi (Figure 1C, “X2”, Late Asturian age). stratigraphic aspects (thickness) in the study of re- The slab is split into two pieces (Plate I, Figs A, constructing seed-fern plants. B), which are both systematically deconstructed and the fragments are examined under a binocular TAXONOMIC EMENDATION microscope at variable magnifications of up to × 25. Eight compound-synangial compressions OF SPECIMES FROM SYDNEY could be collected (Table 1), ignoring fragments COALFIELD smaller than 1 – 2 mm, where the first seven To avoid perpetuating misinterpreted taxa, we tabulated are of unknown positions. The most offer the following emendations. Included in the complete and best preserved eighth one, however, concept of P. krisiae are now the previously originated from just below the surface marked “1a, named: (1) Dolerotheca sp. Zodrow (2007, figs 1b” in Fig. A on Plate I, i.e., part/counterpart 15B-D), and (2) Bernaultia-like sp. (ex Dolerotheca Figs. G and H on Plate II, respectively. Halle), Cleal, Zodrow & Mastalerz (2010, pl. 1, Intact sporangia are freed from the compression figs 6 and 7). (Plate II, Fig. H) in 10 – 15 min treatment with Figure 1. Location map. (A) Canada. (B) The Maritimes Basin. (C) Coal stratigraphy, Sydney Coalfield (Bell 1938, fig. 1), with sample locations of 05-Lst#0 (“X1”) and 08-10/15-9 (“X2”). See also Tables 2 and 1, respectively. 14 E. L. Zodrow, J. Pšenička & Wei-Ming Zhou, Reproductive biology of the arborescent seed-fern Linopteris obliqua 25% v/v hydrofluoric acid, then macerated in the systematic destruction of a 1.7 × 1.3 × 0.46 m Schulze’s solutions maximally for 30 h. Although argillaceous-shaley block, i.e., accession number sporangial cuticles are not observed, the well-pre- 05-Lst#0 in Table 2; see Zodrow et al. (2007, served microspores mounted on glass slides are fig. 23). It originated from an unknown strati- examined at × 500 magnification using a binocu- graphical level in the roof rocks of the Lloyd Cove lar (biological) microscope equipped with Nomarski Seam (Figure 1C, “X1”, Early Cantabrian age). phase-contrast illumination and a digital camera The slab of Fig. A, B on Plate I with its studied for high-resolution images. microspore organs is secured by Erwin L. Zodrow The reference in this paper to Hexagonocarpus sp. (ELZ), and the organ specimens are curated at the is based on the many isolated samples retrieved from West Bohemian Museum, Pilsen, Czech Republic. Table 1. Compound-synangial specimens cited/studied, Sydney Coalfield, Nova Scotia: n.a. not applicable, n.i. not illustrated. Accession no. Figure Analyzed Compression (comp) Preserved size (mm) Impression (imp) Width Height 08-10/15-9-3 Plate II, A n.a. comp 3 10 08-10/15-9-5 Plate II, B n.a. comp 20 10 08-10/15-9-6 Plate II, C n.a. comp 8 10 08-10/15-9-10 Plate II, D n.a. comp 6 7 08-10/15-9-9 Plate II, E n.a. comp 3 4 08-10/15-9-9 n.i. n.a. comp 2 4 08-10/15-9-5 n.i. n.a. comp 2 6 Part and counterpart: 08-10/15-9-1a Plate II, G yes imp 10 17 08-10/15-9-1b Plate II, H yes comp 9 15 Table 2. Summary of Linopteris obliqua pinnules in physical association with Potoniea krisiae or Hexagonocarpus sp. in Sydney Coalfield. *Bedding planes with pinnules. **Pšenička et al. (2019). ***Zodrow et al. (2007, p. 60; 70-72), Hex. = Hexagonocarpus. Accession Coal Seam No. of Sediment Biology of layers No. (Fig.) Layers* Separation L. obliqua P. krisiae Hex. Previous data: 003-252 Roof, 1 m to Single n.a. some 6 n.a. 257 (2) and Lloyd Cove Specimen 004-Z42(2D)** 003-325(3)** Roof, 1 m Single n.a. some 1 n.a. Lloyd Cove Specimen 05-Lst#0 Roof, 1-2 m 4? 4 cm very dense rare none (3)*** Lloyd Cove accumulation or very dense none ca. 80 accumulation Present data: 08-10/15-9 Roof, Hub 2-3 1.5-2 cm very dense 8 none (Plate II, A-H) accumulation Totals 15 > 80 15 Folia vol. 54, No. 1–2, 2020 RESULTS lateral, showing the longitudinal groves; and (3) Plate II, Figs G – I, ventral. Figure C on Plate II Linopteris obliqua, state of occurrences clearly shows an eccentrically positioned, long, and associations narrow (c. 0.3 mm) stalk. All specimens in Plate II The argillaceous-shaley slab (Plate I, Figs A, B) is are decidedly characterized by synangia that fine-grained with dispersed sideritic cementation, extend over the ventral campanulary margins. millimeter bands of siderite, and scattered nodules Remnants of the cap or shoulder ringing the dor- (5 – 20 mm) of highly variables shapes (compare sal portion of, as defined by Pšenička et al. (2019, with Dolerotheca sp. in a sideritic nodule, Zodrow fig. 3A, “PC”) for P. krisiae, are present in Plate & Mastalerz 2019, fig. 2). The pristine preserva- II, Figs B – F, where the thick proximal cap is tion condition of the L. obliqua pinnules and the most clearly preserved in Plate II, Fig. B, arrowed. compound-synangial structures after 12 years of The impression (Plate II, Fig. G) and the com- storage is correlated with the absence of pyrite and pression (Plate II, Fig. H), document the hereto- its destructive oxidative reaction on the compres- fore unknown part/counterpart preservation sions.
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