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Pollen Morphology and Ultrastructure of Quercus With This article was downloaded by: [Paleontological Institute of the Russian Academy of Sciences] On: 14 October 2014, At: 04:11 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Grana Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/sgra20 Pollen morphology and ultrastructure of Quercus with focus on Group Ilex (=Quercus subgenus Heterobalanus (Oerst.) Menitsky): implications for oak systematics and evolution Thomas Denka & Maria V. Teklevab a Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden b A. A. Borissiak Paleontological Institute, Russian Academy of Sciences, Moscow, Russia Published online: 18 Aug 2014. To cite this article: Thomas Denk & Maria V. Tekleva (2014) Pollen morphology and ultrastructure of Quercus with focus on Group Ilex (=Quercus subgenus Heterobalanus (Oerst.) Menitsky): implications for oak systematics and evolution, Grana, 53:4, 255-282, DOI: 10.1080/00173134.2014.918647 To link to this article: http://dx.doi.org/10.1080/00173134.2014.918647 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http:// www.tandfonline.com/page/terms-and-conditions Grana, 2014 Vol. 53, No. 4, 255–282, http://dx.doi.org/10.1080/00173134.2014.918647 Pollen morphology and ultrastructure of Quercus with focus on Group Ilex (=Quercus subgenus Heterobalanus (Oerst.) Menitsky): implications for oak systematics and evolution THOMAS DENK1 & MARIA V. TEKLEVA2 1Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden, 2A. A. Borissiak Paleontological Institute, Russian Academy of Sciences, Moscow, Russia Abstract It has been suggested that pollen ornamentation can be used to distinguish infrageneric groups in Quercus. Here, we document pollen morphology of nearly all species of Quercus Group Ilex and show that they share distinctive microrugulate pollen ornamentation. Furthermore, pollen ultrastructure of all six infrageneric groups of Quercus was studied comparatively and evaluated using a phylogenetic framework for Fagaceae. Characteristics of the foot layer, such as its thickness and continuity, and the ratio foot layer to tectum are not stable within infrageneric groups of Quercus. The foot layer in derived groups (Groups Cerris, Quercus, Lobatae) is irregular, discontinuous and thin (type 2 foot layer). Members of Groups Cyclobalanopsis and Ilex either have a regular, continuous, and comparatively thick foot layer (type 1), or a type 2 foot layer. Only Group Protobalanus consistently has a type 1 foot layer and is similar to Trigonobalanus and Colombobalanus among trigonobalanoids, and to castanoids. Because (I) all except for one species of Castanea and Nothofagus have a type 1 foot layer and (II) basal groups in Quercus may have a type 1 foot layer, we suggest that this is the ancestral state in Fagaceae, and that the type 2 foot layer of Fagus is derived. Keywords: Quercus Group Ilex, evolution, pollen ornamentation, foot layer A major attempt of phylogenetic and evolutionary Denk 2003; Huysmans et al. 2003; Dettmann & studies has been to explore morphological characters Jarzen 2005; Denk & Tekleva 2006; Wortley et al. or combinations of characters that are suitable to 2007, 2008; Denk & Grimm 2009; Nagels et al. define natural groups of plants (Walker & Doyle 2009; Wang et al. 2010). 1975; Doyle & Endress 2000, 2010; Doyle 2005). Quercus is a large genus (c. 400–500 species) of Existing phylogenies for a large number of plant monoecious trees and shrubs distributed across the groups based on molecular data can provide a frame- Northern Hemisphere with a small extension into work for assessing the evolution and phylogenetic South America. Six infrageneric groups are recog- significance of morphological characters (see e.g. nised within Quercus: (I) Group Cyclobalanopsis (c. Doyle & Endress 2000; Endress & Doyle 2009). At 150 spp.; estimated by Flora of China Editorial the same time, conflict in molecular phylogenetic Committee 1999) occurs in subtropical and tropical Downloaded by [Paleontological Institute of the Russian Academy Sciences] at 04:11 14 October 2014 reconstructions can be investigated using morpholo- regions in southeast Asia, while the Groups Lobatae gical characters (Denk et al. 2005; Friis et al. 2007; (II) (c. 195 spp.; estimated by Flora of North America Denk & Grimm 2009; Mathews 2009). Pollen char- Editorial Committee 1997) and Protobalanus (III) acters have been playing a key role in identifying (five species) at present are found in the New World natural groups and in understanding plant evolution only. The Groups Cerris (IV) and Ilex (V) (11 and c. (e.g. Kuprianova 1962; Doyle & Le Thomas 1997; 30 spp.; Denk & Grimm 2009, 2010) are confined to Correspondence: Thomas Denk, Palaeobiology Department, Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm, Sweden. E-mail: [email protected] (Received 8 January 2014; accepted 25 March 2014) © 2014 Collegium Palynologicum Scandinavicum 256 T. Denk and M. V. Tekleva Eurasia, while the Group Quercus (VI) (c. 160 spp.; first, to document the variability of the tectum estimated in Flora of North America Editorial ornamentation within and among species of Committee 1997 and Flora of China Editorial Quercus Group Ilex, and second, to evaluate the Committee 1999) occurs both in the Old and New evolutionary significance of pollen ultrastructure World. The majority of molecular phylogenetic stu- encountered in members of all six infrageneric dies have resolved two major clades within Quercus, groups of Quercus and its relevance for Fagaceae one consisting of the infrageneric groups Ilex, Cerris evolution. and Cyclobalanopsis, and the other consisting of the groups Protobalanus, Lobatae and Quercus (Manos & Stanford 2001; Manos et al. 2001, 2008;Oh& Material and methods Manos 2008; Denk & Grimm 2010). In traditional classification schemes of Quercus (e.g. Pollen material investigated for the present study Camus 1936–1938, 1938–1939, 1952–1954; Schwarz (Table I) originates from herbarium material stored 1936;Nixon1993), the systematic placement of at the herbaria Edinburgh (E), Istanbul (ISTO), Groups Cerris and Ilex (the ‘Cerris oaks’ and the Stockholm (S) and Berkeley (UC). LM investiga- ‘Ilex oaks’) has been remarkably unstable (cf. Denk & tions were made on material from the palynological Grimm 2010 for the taxonomic history of these slide collection at the Swedish Museum of Natural groups) because of the lack of diagnostic morphologi- History, Stockholm. A full list of herbarium vou- cal characters. chers for LM, SEM and TEM preparations is pro- Recently, it has been shown that pollen orna- videdinthe‘Specimens investigated’ section. For mentation is highly diagnostic for the six infrage- SEM investigations, pollen was removed from neric groups of Quercus (Denk & Grimm 2009). anthers and mounted on SEM stubs. Non-aceto- Furthermore, it has earlier been suggested that lysed pollen was used for SEM observations, Ilex oaks are distinctive among oaks by having a because previous studies using acetolysed and relatively thicker and more homogeneous foot non-acetolysed pollen of Fagaceae showed the layer than other groups of oaks (Crepet & same details of the pollen surface (cf. Praglowski Daghlian 1980). The latter study was based on 1982, 1984; Solomon 1983a, 1983b;Denk2003; very limited material (two specimens of Quercus Denk & Grimm 2009). Group Ilex). For TEM investigations, individual anthers or parts At present, there is no comprehensive study doc- of anthers were fixed with 1% osmium tetroxide umenting pollen morphology in the European and (OsO4), dehydrated in an ethanol series, stained with western and eastern Asian members of Quercus uranyl acetate, dehydrated in acetone, and embedded Group Ilex. A number of detailed accounts on pollen in epoxy resin. Pollen grains were sectioned with an morphology and ultrastructure of genera of Fagales ultramicrotome LKB-3 and ultrathin sections were are available (Alfaroa, Carya, Castanea, Castanopsis, then post-stained with lead citrate, and examined Chrysolepis, Engelhardia, Fagus, Juglans, Lithocarpus, using a Jeol 100 B and a Jeol 1011 transmission elec- Nothofagus, Oreomunnea, Platycarya, Pterocarya; tron microscopes.
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