Seagrass Foliage from the Maastrichtian Type Area (Maastrichtian, Danian, NE Belgium, SE Netherlands) ⁎ R.W.J.M
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Review of Palaeobotany and Palynology 144 (2007) 301–321 www.elsevier.com/locate/revpalbo Seagrass foliage from the Maastrichtian type area (Maastrichtian, Danian, NE Belgium, SE Netherlands) ⁎ R.W.J.M. van der Ham a, , J.H.A. van Konijnenburg-van Cittert a,b, L. Indeherberge c a Nationaal Herbarium Nederland, Leiden branch, P.O. Box 9514, 2300 RA Leiden, The Netherlands b Nationaal Natuurhistorisch Museum, P.O. Box 9517, 2300 RA Leiden, The Netherlands c Reuvoortweg 63, 3520 Zonhoven, Belgium Received 17 November 2005; received in revised form 3 July 2006; accepted 5 July 2006 Available online 18 September 2006 Abstract Seagrasses are the only angiosperms that have invaded the marine environment. Fossil seagrasses are rare and received only little attention to date. However, among the scarce plant fossils in the marine deposits of the Maastrichtian type area, remains of seagrasses are relatively common. The present paper provides a detailed description of the morphology and anatomy of the leaves. Maastrichtian and Danian material is considered here as a single species: Thalassotaenia debeyi gen. nov., sp. nov. Campanian material is assigned to Zosterites, but this is not necessarily a seagrass. Thalassotaenia includes the oldest anatomically preserved material of seagrass leaves. Comparison with extant seagrasses shows that there is not an easy match with any of the extant genera or families. Amphibolis and Thalassodendron (Cymodoceaceae) and Posidonia (Posidoniaceae) show the greatest resemblance. Possibly, Thalassotaenia represents the leaves of an ancestor of the Cymodoceaceae–Posidoniaceae (–Ruppiaceae) clade found in several phylogenetic analyses. Because Thalassotaenia also shows some features not found in extant seagrasses, it might also represent an extinct clade of seagrasses. © 2006 Elsevier B.V. All rights reserved. Keywords: seagrass leaves; Cymodoceaceae; Posidoniaceae; Thalassocharis; Maastrichtian 1. Introduction tidae (Monocotyledonae) having been recognised in a rbcL study by Les et al. (1997). Considerable monocot diver- Worldwide, seagrass communities are important eco- sification took place during the Early Cretaceous, and systems in relatively warm, shallow seas (Green and Short, seagrasses probably evolved from freshwater monocots 2003). They are well-known for their high biodiversity and during the Late Cretaceous (Janssen and Bremer, 2004). over the last decades there has been a lot of research into Fossil seagrasses are rare. The oldest record is from the many aspects of seagrass biology and ecology (e.g. Brasier, Early Campanian of the Netherlands: Thalassocharis 1975; Larkum et al., 1989, 2006). Seagrasses are the only mülleri (Debey, 1848; Pomel, 1849; Debey,1851), which angiosperms that have successfully invaded the marine represents the stem of a seagrass. Voigt and Domke (1955) environment. They make up an ecological, polyphyletic published an extensive paper on Thalassocharis bosquetii group rather than a single taxonomical, monophyletic Debey ex Miquel from the Late Maastrichtian of the group, three different clades within the subclass Alisma- Netherlands, in which they analysed anatomically pre- served stems and attached roots, considering these related ⁎ Corresponding author. Fax: +31 0 71 527 35 11. to those of the extant genus Cymodocea. A description of E-mail address: [email protected] (R.W.J.M. van der Ham). the leaves is lacking so far. They are always found detached 0034-6667/$ - see front matter © 2006 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2006.07.008 302 R.W.J.M. van der Ham et al. / Review of Palaeobotany and Palynology 144 (2007) 301–321 and incomplete, without base and apex, represented by different parts of the Maastrichtian type area (Fig. 1) fragments mostly not longer than a few centimeters, and and originates from the Early Campanian to Early usually poorly preserved, showing only a faint imprint with Danian time interval (Fig. 2). indistinct venation. Recently, however, several leaf frag- Two Early Campanian specimens are from the Vaals ments with remarkable details were discovered by one of us Formation (Inoceramus lingua/Gonioteuthis quadrata (LI). These include isolated fragments in chert, showing Zone) of Hallembaye in the southwestern part of the venation, fiber strands and even cell patterns, as well as the area (Fig. 1: locality 1). remains of amassed fragments in chert suited for thin- A single Late Maastrichtian specimen is from the Lixhe sectioning. This material allows detailed description and I Member of the Gulpen Formation (Belemnitella junior statements about the affinity of the seagrass foliage from Zone) of Loën in the southwestern part (Fig. 1: locality 2), the Maastrichtian type area (NE Belgium, SE Netherlands). while all other Late Maastrichtian material is from several The present paper is part of the ongoing study of members of the Maastricht Formation (Belemnitella junior fossil plants from the Maastrichtian type area. Earlier Zone and Belemnella kasimiroviensis Zone) in both the papers dealt with conifers and associated fungi (Van der southwestern and the northeastern part of the area. The Ham and Dortangs, 2005, and references therein). material from the southwestern part (Fig. 1: localities 1, 3– 6) is from the fully marine facies of the Maastricht 2. Material, localities and stratigraphy Formation (Valkenburg to Meerssen Members), while specimens from the northeastern part (Fig. 1:localities8– The material studied (c. 200 specimens; see Section 20) originate from the Kunrade Chalk, which is the more 4. Systematics and descriptions) was collected in coastal facies of the Maastricht Formation (Fig. 2). The Fig. 1. Map of the Maastrichtian type area, showing the localities where Thalassotaenia debeyi has been collected: 1. Hallembaye, 2. Loën, 3. Sluizen, 4. Romontbos, 5. Zichen, 6. ENCI, 7. Geulhem, 8. boring XVI, 9. boring S.M. XIX, 10. Emma I, 11. Oranje Nassau III, 12. boring 634, 13. boring 1291, 14. Kunrade, 15. Winthagen, 16. Welterberg, 17. Daelhoeve, 18. Benzenrade, 19. Simpelveld, 20. Vetschau. Locality 1 (Hallembaye) also provided the material of Zosterites sp. R.W.J.M. van der Ham et al. / Review of Palaeobotany and Palynology 144 (2007) 301–321 303 Fig. 2. Stratigraphy of the Cretaceous deposits in the Maastrichtian type area, showing the origin of the material of Thalassotaenia debeyi (Gulpen and Maastricht Formations) and Zosterites sp. (Vaals Formation). See Jagt (1999) for further stratigraphical details. The dot at the level of the Lichtenberg Horizon represents the specimens from the residual chert deposit (′flint eluvium′) of Hallembaye (including the type material) as well as those from the Valkenburg Member, and the dot at the Gronsveld Member, besides those of the Gronsveld Member, also the specimens from the Kunrade Chalk. Besides the residual chert deposit derived from the top of the Lanaye Member and base of the Maastricht Formation (indicated in scheme as ′flint eluvium′), a younger residual chert deposit exists (not indicated in scheme) derived from a part of the Nekum Member. 304 R.W.J.M. van der Ham et al. / Review of Palaeobotany and Palynology 144 (2007) 301–321 Fig. 3. Selected specimens of Thalassotaenia debeyi, gen. et sp. nov., showing veins, alternating fiber strands and cross-veins. a–c. Kunrade Chalk, Emma I, a. JMS 52006 (see also Plate III, 3), b, c. JMS 52067 (see also Plate III, 1). d–e. Residual chert deposit (′flint eluvium′), Hallembaye, NHMM LI 451 and 452, respectively (part and counterpart; type material; see also Plate I, 3, 4); note adaxial and abaxial fiber strands close together in Fig. 3d, immediately right of the midrib. f–g. Kunrade Chalk, Benzenrade, f. JMS 52027 (see also Plate II, 2), g. JMS 52045. h. Nekum Member, residual chert deposit, Sluizen, NHMM LI 4606 (see also Plate I, 2). i. Early Danian part of Meerssen Member, Geulhem, NHMM RH 808. Bar=10 mm (a–e, g–i) or 20 mm (f ). R.W.J.M. van der Ham et al. / Review of Palaeobotany and Palynology 144 (2007) 301–321 305 anatomically preserved leaves were found in residual chert vein thicker than the others; stomata absent; hypoder- deposits (‘flint eluvium’), which originated by dissolution mis present. of the chalky matrix from the top of the Lanaye Member of Etymology: From Greek thalassa (sea) and tainia the Gulpen Formation and the base of the Maastricht (strap), referring to the shape of the leaves and the Formation (Hallembaye; Fig. 1: locality 1) and from the environment in which they grew. Nekum Member (Romontbos quarry and Sluizen; Fig. 1: Type: Thalassotaenia debeyi sp. nov. localities 3, 4). Affinity: Monocotyledonae, ?Cymodoceaceae/Posido- Early Danian fossils are from the uppermost part of niaceae (see: Section 6. Affinities). the Meerssen Member of the Maastricht Formation (E clay sensu Smit and Brinkhuis, 1996) of Geulhem Thalassotaenia debeyi Van der Ham et Van Konij- (Fig. 1: locality 7). nenburg-Van Cittert, sp. nov. (Fig. 3; Plates I, 2–4; II; The following abbreviations are used to denote the III; IV, 1–3; V, 1–3;) locations of the specimens: IRSNB: Institut royal des Sciences naturelles de Belgique, Brussels, Belgium; JMS: 1849 Abdrücke eines najadeenartigen Blattes — Debey, p. 285, 286 Jongmans collection, Nationaal Natuurhistorisch Muse- 1851 Najadeenblätter (Thalassocharis/Zosterites) — Debey, p. 568 um, Leiden, The Netherlands; MB.PB.: Museum für 1853 ? phyllitae monocotylei,1.— Miquel, p. 53 Naturkunde der Humboldt Universität, Berlin, Germany; 1853 non phyllitae monocotylei,2.— Miquel,