Cordaites Principalis (Germer) Geinitz, Carboniferous Maritimes Basin, Canada
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Functional groups and elemental analyses of cuticular morphotypes of Cordaites principalis (Germer) Geinitz, Carboniferous Maritimes Basin, Canada Zodrow, Erwin L; Mastalerz, Maria; Orem, William H; Šimunek, Zbynek; Bashforth, Arden Roy Published in: International Journal of Coal Geology Publication date: 2000 Document version Publisher's PDF, also known as Version of record Citation for published version (APA): Zodrow, E. L., Mastalerz, M., Orem, W. H., Šimunek, Z., & Bashforth, A. R. (2000). Functional groups and elemental analyses of cuticular morphotypes of Cordaites principalis (Germer) Geinitz, Carboniferous Maritimes Basin, Canada. International Journal of Coal Geology, 45(1), 1-19. Download date: 24. sep.. 2021 International Journal of Coal Geology 45Ž. 2000 1±19 www.elsevier.nlrlocaterijcoalgeo Functional groups and elemental analyses of cuticular morphotypes of Cordaites principalis ž/Germar Geinitz, Carboniferous Maritimes Basin, Canada Erwin L. Zodrowa,) , Maria Mastalerzbc , William H. Orem , ZbynekÆÊ SimunekÆ d , Arden R. Bashforth e,1 a Department of Earth Sciences, UniÕersity College of Cape Breton, Sydney, NoÕa Scotia, Canada B1P 6L2 b Indiana Geological SurÕey, Indiana UniÕersity, 611 N. Walnut GroÕe, Bloomington, IN 47405-2208, USA c U.S. Geological SurÕey, MS 956, National Center, Reston, VA 22092, USA d Czech Geological SurÕey, KlaroÂÕ 3r131, 118 21 Praha 1, Czech Republic e Department of Earth Sciences, Memorial UniÕersity of Newfoundland, St. John's, Canada, A1B 3X5 Received 29 July 1999; accepted 19 June 2000 Abstract Well-preserved cuticles were isolated from Cordaites principalis Ž.Germar Geinitz leaf compressions, i.e., foliage from extinct gymnosperm trees Coniferophyta: Order Cordaitales. The specimens were collected from the Sydney, Stellarton and Bay St. George subbasins of the once extensive Carboniferous Maritimes Basin of Atlantic Canada. Fourier transformation of infrared spectraŽ. FTIR and elemental analyses indicate that the ca. 300±306-million-year-old fossil cuticles share many of the functional groups observed in modern cuticles. The similarities of the functional groups in each of the three cuticular morphotypes studied support the inclusion into a single cordaite-leaf taxon, i.e., C. principalis Ž.Germar , confirming previous morphological investigations. Vitrinite reflectance measurements on coal seams in close proximity to the fossil-bearing sediments reveal that the Bay St. George sample site has the lowest thermal maturity, whereas the sites in Sydney and Stellarton are more mature. IR absorption and elemental analyses of the cordaite compressions corroborate this trend, which suggests that the coalified mesophyll in the leaves follows a maturation path similar to that of vitrinite. Comparison of functional groups of the cordaite cuticles with those from certain pteridosperms previously studied from the Sydney Subbasin shows that in the cordaite cuticles highly conjugated C±OŽ 1632 cmy1. bands dominate over carbonyl stretch that characterizes the pteridosperm cuticles. The differences demonstrate the potential of chemotaxonomy as a valuable tool to assist distinguishing between Carboniferous plant±fossil groups. Published by Elsevier Science B.V. Keywords: Cordaites compression leaves; cuticles; Carboniferous; FTIR; elemental analyses ) Corresponding author. 503 Coxhead Road, Sydney, N.S., Canada B1R 1S1. E-mail addresses: [email protected]Ž. E.L. Zodrow , [email protected] Ž M. Mastalerz . , [email protected] Ž. W.H. Orem , [email protected]Ž. Z. SimunekÆ Ê , [email protected] Ž A.R. Bashforth . 1 Presently at Anderson Exploration, 324 8th Ave. SW Calgary, Alberta, Canada T2P 2Z5. 0009-2541r00r$ - see front matter. Published by Elsevier Science B.V. PII: S0166-5162Ž. 00 00018-5 2 E.L. Zodrow et al.rInternational Journal of Coal Geology 45() 2000 1±19 1. Introduction Ž.Fig. 1B and C . This circumstance provided the significant opportunity for a comprehensive study of Cordaitean fossil leaves Ž.Cordaites, Unger 1850 the cordaitean compressions in the basin. As a result, are known from early Carboniferous to early Per- five new cuticular morphotypes were established for mian deposits of the Euramerican palaeokingdom the Carboniferous leaf taxon Cordaites principalis Ž.Rothwell, 1988 , representing a lifespan of ca. 80 Ž.Germar Geinitz Ž Zodrow et al., 2000 . million years. The cordaites trees from which the The impetus for the present study stems from the cordaitean leaves originated constituted a diverse recognition that future progress in phylogeny of Eu- group of extinct gymnospermous plants that had ramerican cordaiteans requires more adequate taxo- extremely variable growth habits and palaecological nomic parameters than are presently available. In this tolerances. Mire-dwelling cordaites trees included investigation, methods of Fourier transformation of mangrove-like forms up to 5 m tall with aerenchy- infrared spectraŽ. FTIR and elemental analyses are matous, stilt-like rootsŽ Cridland, 1964; Raymond, applied to three representatives of the five cordaitean 1988; Raymond and Phillips, 1983; Costanza, 1985. , morphotypes. The purpose is to assess the applicabil- and small, thicket-forming, understory shrubs ity of these techniques in classification, and to deter- Ž.Rothwell and Warner, 1984; Costanza 1985 . By mine which parameters could be of importance for comparison, cordaites trees growing on drier, well- future research in cordaitean chemotaxonomy. drained clastic substrates, such as floodplains, river levees, or extrabasinal lowlands were very tall, for- est-forming trees that towered tens of meters high ŽGrand'Eury, 1877; Mapes and Gastaldo, 1986; 2. Methods Rothwell, 1988; Rothwell and Mapes, 1988. Cordaitean leaves, which can reach 1 m in length, Cuticles were prepared by macerating the com- are strap-like or linear in shape with longitudinal pressions from 7 h to 3 days in Schulze's oxidizing venation pattern that parallels the leaf marginŽ Fig. solution that consisted of 150 ml 70% nitric acid 1A. The foliage is a common component of com- with 3±5 g dissolved potassium chlorate. After oxi- pression assemblages and has been exhaustively in- dation, samples were neutralized in a 4.5% ammo- vestigated since Grand'EuryŽ. 1877 . Based on com- nium hydroxide solution, and well rinsed in deminer- parative cuticle morphology, FlorinŽ. 1931, 1951 alized water. Through this process, 200 cuticular postulated that cordaites were ancestral to conifer mounts were prepared, and five new cuticular mor- trees, but the exact nature of the relationship between photypes of C. principalis Ž.Germar were erected the two taxa is equivocalŽ Meyen, 1984; Crane 1985; Ž.Zodrow et al., 2000 . Representative samples of Rothwell, 1986. Alternatively, both taxa may have three of these cuticular morphotypes and their com- been derived from a common progenitorŽ Rothwell, pressions were investigated. Compression mounts 1977. , such as the progymnospermous Ar- were routinely examined under a light microscope to chaeopteris Ž.Beck, 1981 , or a pteridosperm Ž Roth- determine the presence of pyrite and other minerals, well, 1988. and to observe variation in coalification patterns. Cordaitean compressions were first recorded from Specimens for FTIR were prepared using the the Carboniferous in New Brunswick and Nova Sco- potassium bromideŽ. KBr pellet technique. A very tia, Canada, by DawsonŽ. 1868, 1891 , and studied small amount of the cuticle or compressionŽ ap- systematically by BellŽ. 1940, 1944, 1962 . Since the proximately 2 wt.% of the mixture. was mixed with fossil-bearing strata in the Carboniferous Maritimes finely ground KBr to produce pellets. These were Basin covering parts of Nova Scotia, New Brunswick, analyzed on a Nicolet 20SXC spectrometer, equipped and NewfoundlandŽ. Fig. 2 generally have lower with a DTGS detector, at a resolution of 4 cmy1, thermal maturity, as a result of having experienced collecting 1024 scansrsample. The infrared signal limited tectonism and burialŽ Hacquebard and Ž.IR was recorded in the region between 400 and Cameron, 1989. , cordaitean cuticles are well pre- 4000 cmy1 wavenumber. Bands were identified by served and show delicate stomatalrepidermal details comparison with published assignmentsŽ Painter et E.L. Zodrow et al.rInternational Journal of Coal Geology 45() 2000 1±19 3 Fig. 1. Cordaites principalis Ž.Germar Geinitz, compressions and cuticles. Ž. A Fragments of leaf compressions with typically thick, primary veins parallel to the foliar margins. Bay St. George Subbasin, Newfoundland. B-24, cm scale.Ž. B Intact adaxial Ž. ad and abaxial Ž. ab cuticle, showing the typically elongate cells for morphotype 1, Bay St. George Subbasin, B-228. Scale bars60 mm.Ž. C Intact abaxial cuticle of morphotype 4, showing stomatal apparatusŽ. o . Sydney Subbasin, Nova Scotia, SY-211. Scale bars100 mm. Ž D . Mesophyllous matter on an inner surface of a cuticle from Bay St. George Subbasin. Scale bars10 mm. al., 1981; Wang and Griffith, 1985; Sobkowiak and Elemental analyses for C, H, N, O, and S in the Painter, 1992. compressions and morphotypes were performed on 4 E.L. Zodrow et al.rInternational Journal of Coal Geology 45() 2000 1±19 Fig. 2. Carboniferous Maritimes Basin of eastern, Canada, and study areas of the three subbasins. dried samplesŽ. 608C on a Carlo Erba Elemental ous Maritimes BasinŽ. Fig. 2 . In the Sydney Sub- AnalyzerŽ. Lyons et al., 1995 . Mineral matter is basin, the compressions are entombed in grey, fis- defined as the difference between the summed