ebook img

Early Evolution of Flowers PDF

230 Pages·1994·9.84 MB·English
Save to my drive
Quick download
Download
Most books are stored in the elastic cloud where traffic is expensive. For this reason, we have a limit on daily download.

Preview Early Evolution of Flowers

Plant Systematics and Evolution . Supplement 8 P. K. Endress and E. M. Friis (eds.) Early Evolution of Flowers Springer-Verlag Wien New York Prof. Dr. PETER K. ENDRESS Institut fur Systematische Botanik der Universitat, Zurich, Schweiz Prof. Dr. ELSE MARIE FRIIS Naturhistorika Riksmuseet Sektionen for Paleobotanik, Stockholm, Sweden This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machines or similar means, and storage in data banks. © 1994 Springer-Verlag/Wien Softcover reprint ofthe hardcover 1st edition 1994 Typesetting: Thomson Press (India) Ltd., New Delhi, 110001 Printed on acid-free and chlorine-free bleached paper With 130 Figures Library of Congress Cataloging-in-Publication Data Early evolution of flowers/Po K. Endress, E. M. Friis (eds.). p. cm. - (Plant systematics and evolution. Supplement; 8) ISBN-13: 978-3-7091-7432-6 e-ISBN-13: 978-3-7091-6910-0 DOl: 10.1007/978-3-7091-6910-0 1. Flowers. 2. Angiosperms - Evolution. 3. Flowers - Morphology. I. Endress, Peter K. II. Friis, Else Marie. III. Series: Plant systematics and evolution. Supplementum; 8. QK653.E2 1994 582.13'04463 - dc20 ISSN 0172-6668 Contents ENDRESS, P. K., FRIIS, E. M.: Introduction-Major trends in the study of early flower evolution DOYLE, J. A.: Origin of the angiosperm flower: a phylogenetic perspective 7 FRIIS, E. M., PEDERSEN, K. RAUNSGAARD, CRANE, P. R.: Angiosperm floral structures from the Early Cretaceous of Portugal 31 CRANE, P. R., FRIIS, E. M., PEDERSEN, K. RAUNSGAARD. Palaeobotanical evidence on the early radiation of magnoliid angiosperms 51 CREPET, W. L., NIXON, K. c.: Flowers of Turonian Magnoliidae and their implications 73 DRINNAN, A. N., CRANE, P. R., HOOT, S. B.: Patterns of floral evolution in the early diversification of non-magnoliid dicotyledons (eudicots) 93 NISHIDA, H.: Elsemaria, a Late Cretaceous angiosperm fructification from Hokkaido, Japan 123 TUCKER, S. c., BOURLAND, J. A.: Ontogeny of staminate and carpellate flowers of Schisandra glabra (Schisandraceae) 137 WILLIAMSON, P. S., SCHNEIDER, E. L.: Floral aspects of Barclaya (Nymphaeaceae): pollination, ontogeny and structure 159 ENDRESS, P. K.: Evolutionary aspects of the floral structure in Ceratophyllum 175 KOSUGE, K.: Petal evolution in Ranunculaceae 185 ERBAR, c., LEINS, P.: Flowers in M agnoliidae and the origin of flowers in other subclasses of the angiosperms. I. The relationships between flowers of Magnoliidae and Alismatidae 193 LEINS, P., ERBAR, c.: Flowers in M agnoliidae and the origin of flowers in other subclasses of the angiosperms. II. The relationships between flowers of Magnoliidae, Dilleniidae, and Caryophyllidae . 209 Taxonomic Index 219 Subject Index 225 Listed in Current Contents Pl. Syst. Evol. [Suppl.] 8: 1-6 (1994) Introduction - Major trends in the study of early flower evolution Our knowledge of early angiosperm phylogeny and of flower evolution is rapidly expanding. New evidence comes from different directions, and it seems important to constantly synthesize new results from these fields. (1) Palaeobotany brought two major developments, first comparative structural and stratigraphic pollen studies (since BRENNER 1963 and DOYLE 1969), followed by the discovery of excellently preserved Cretaceous flowers (since FRIIS & SKARBY 1981). (2) Development and biology of extant flowers and structure of modern pollen was studied in many living conservative groups during the same period. Application of the SEM for comparative floral development was especially helpful (since ENDRESS 1972). The comparative study of pollen with combined SEM and TEM techniques provided a powerful approach to link extant and fossil material (since WALKER 1976). Comparative pollination biology of primitive flowers in relation to floral structure brought another new dimension (since GOTTSBERGER 1974). (3) The application of cladistic techniques facilitated the rigorous discussion on the origin of the angio sperms from other seed plants and on early angiosperm diversification (since CRANE 1985, DOYLE & DONOGHUE 1986). (4) More recently comparative molecular studies with DNA and RNA became possible due to new techniques, applied for macro systematics of angiosperms (since JANSEN & PALMER 1987). (5) The latest develop ment is the direct analysis of floral development with molecular genetic techniques, which, however, as yet has been done only for few model species of evolutionarily more advanced angiosperm groups (COEN & MEYEROWITZ 1991). Results from the fossil record The fossil record of pollen and leaves has provided a rough framework for under standing patterns in early angiosperm evolution (BRENNER 1963, 1976; DOYLE & HICKEY 1976; HUGHES 1976). Based on these organs DOYLE & HICKEY (1976) demonstrated a progressive complexity and diversity in angiosperm morphology in the Early to mid-Cretaceous Potomac group sequence from eastern North America. Recent discoveries of well-preserved angiosperm floral structures from Cretaceous strata in Europe and North America indicate that a parallel increase in complexity in floral organs took place during the same time interval (e.g., FRIIS & CREPET 1987, FRIIS & ENDRESS 1990). Although the fossil floral structures are much more rare in the Cretaceous strata than are leaves and pollen, they provide a broader and more profound basis for comparison with modern angiosperms and the establishment of systematic affinities of early angiosperms. Results obtained 2 P. K. Endress & E. M. Friis: so far from the study of Cretaceous flowers support the generally accepted view that the magnoliids contain the most primitive angiosperms, and that the hamame lidids diversified from the magnoliids early in the history of angiosperms. Early Cretaceous flowers are typically minute, unisexual or bisexual with few floral parts, undifferentiated perianth, massive stamens typically with valvate dehiscence, and uniovulate carpels, indicating that flowers of generallauralean or piperalean organi zation were well-established in the Early Cretaceous (e.g., FRIIS & ENDRESS 1990, CRANE & al., contribution 3 in this volume). Larger floral structures related to the woody M agnoliales are rare in the Cretaceous fossil record (DILCHER & CRANE 1985) and appeared later than the smaller flowers. The abundance of small simple flowers early in the history of angiosperms may reflect a particular rapid diversification of lauralean taxa rather than reflecting the basal floral organization in angiosperms, and rooting of the angiosperms is still much debated (contribution by DOYLE in this volume). Based on leaf architecture and sedimentological data DOYLE & HICKEY (1976) suggested that early angiosperms were riparian weeds of magnoliid and monocot affinity (paleoherbs). A herbaceous growth form was also suggested by TAYLOR & HICKEY (1990) based on the leaf architecture of a small compression/impression fossil from the Aptian of Australia. Little is known about growth form in early angiosperms, but the scarcity of larger pieces of angiosperm wood in the Cretaceous, particularly in the Early and mid-Cretaceous, could be an indication for the absence of larger angiosperm trees in the early phases of angiosperm diversification (UPCHURCH & WOLFE 1987, HERENDEEN 1991). While the fossil record clearly indicates that the first major radiation of angiosperms took place in the Early Cretaceous (e.g., DOYLE & HICKEY 1976, CRANE & LrDGARD 1989) phylogenetic analysis indicates that angiosperms may have originated much earlier, possibly in the Triassic (DOYLE & DONOGHUE 1993, CRANE 1993). An early radiation of angiosperms has also been claimed by CORNET (e.g., 1989a, b) based on dispersed pollen and the enigmatic Sanmiguelia plant. However, the nature of these fossils is still poorly understood and their angiosperm affinity remains to be established. Although several of the dispersed pollen grains show angiospermous features such as reticulate and tectate exine (e.g., CORNET 1989b), details in wall ultrastructure and aperture configuration indicate that they are more related to non-angiospermous anthophytes (see discussion in FRIIS & PEDERSEN 1995). Results from the study of the structure and biology of extant flowers The extant M agnoliidae contain many families with a single or only few species. In the classification of CRONQUIST (1988) of the 39 families 14 contain a single genus. Some of the families with more than one genus have a pronouncedly scattered relictual distribution (such as Chloranthaceae, Winteraceae, Lardizabalaceae), and some are known to have occurred in a much wider area in the Cretaceous or early Tertiary {e.g., Chloranthaceae, Winteraceae) and have been more diverse than today (KRUTZSCH 1989). Spiral phyllotaxis, which is mostly considered to be the basal pattern of arrangement of floral organs, is less generally common in the extant M agnoliidae Introduction - Major trends in the study of early flower evolution 3 than was earlier believed (TUCKER 1984, ENDRESS 1986). Irregular and whorled patterns are equally common. All these patterns may co-occur at low systematic levels, even within single species. Variability in floral organ number may be equally extensive (ENDRESS 1986). There are many families with predominantly unicarpellate flowers and many with uniovulate carpels. There are several families without a distinct perianth, and many with a simple perianth that is not differentiated into calyx and corolla. The delimitation of a flower towards the bract region outside the flower is not always clear-cut. Valvate dehiscence of anthers is widespread (ENDRESS & HUFFORD 1989). In some groups, especially in Ceratophyllales, Piperales and in some Laurales, the flowers are small, sometimes exceedingly so. Large, beetle-pollinated flowers show peculiar specializations. Their organization does seem less primitive than that of some smaller, less functionally differentiated flowers (GOTTSBERGER 1974, 1988; GOTTSBERGER & al. 1980). Flies or small moths may contain pollinators as primitive as beetles (THIEN & al. 1985, PELLMYR 1992, KATO & INOUE 1994). Floral secretions, such as nectar and stigmatic fluid, in addition to pollen, may constitute important rewards for pollinators in magnoliid flowers. Protogyny is the dominant kind of dichogamy in bisexual flowers (ENDRESS 1990), which could also indicate that secretions, and not pollen, were a primitive reward. Optical and olfactorial attraction of pollinators is by stamens and carpels in many groups, and not by the perianth. The advent of anatropous ovules may have preceded that of closed carpels (TOMLINSON 1991, LLOYD & WELLS 1992). However, some paleo herbs, such as Ceratophyllaceae and Piperales, and also Chloranthaceae, have orthotropous ovules, which may be viewed as apomorphies due to particular ovary architectures. A consequence of these various new developments is that classification of major angiosperm groups is under extensive revision. The application of some informal names for the most problematical groups, such as, e.g., "paleo herbs", seems wise before a new classification can be worked out on a relatively consolidated new basis. As a working basis we here still use the classification by CRONQUIST (1988) in addition to the informal names. As its appears now, earlier assumptions on primitive angiosperms were not completely wrong. What remains valid in particular is that the M agnoliidae are the most basal group. However, it has become uncertain whether the woody M agnoliales are at the base. There are equally good arguments to place herbaceous groups (paleoherbs) at the base of the Magnoliidae (e.g., DOYLE & HICKEY 1976, ZIMMER & al. 1989, LES & al. 1991, TAYLOR & HICKEY 1992, CHASE & al. 1993, DOYLE & DONOGHUE 1993, QIU & al. 1993, and discussion in the contribution by DOYLE in this volume). This volume includes new advances in our understanding of early angiosperm and flower evolution and brings together contributions from palaeobotanical as well as neobotanical studies. The contributions were presented at two symposia held at the XV International Botanical Congress in Yokohama on September 1, 1993, and organized by PETER K. ENDRESS, ELSE MARIE FRIIS, HARUFUMI NISHIDA, and MICHIO TAMURA. The first contribution discusses various hypotheses on the evolution of flowers and floral organs based on recent phylogenetic-cladistic analyses (DOYLE). The 4 P. K. Endress & E. M. Friis: following three contributions give a survey of the vast diversity of Cretaceous magnoliid floral organs discovered during the past few years, and two chapters deal with higher dicots. Early Cretaceous assemblages from Portugal (FRIIS & aI., contribution 2) and from eastern North America (CRANE & aI., contribution 3) exhibit a wide range of magnoliid fossils, and particularly taxa related to the Laurales appear to be well differentiated, but it is characteristic that none of the fossils can be assigned with certainty to a modern family. Magnoliid fossils continue to be important also in Late Cretaceous (Turonian) floras of eastern North America and include several taxa that are comparable to extant taxa at the family level (CREPET & NIXON, contribution 4). The earliest diversification of the Hamamelididae started in the Early Cretaceous; and a large variety of taxa are present by the Late Cretaceous (DRINNAN & aI., contribution 5). By that time most higher dicotyledon subclasses were established, and the new genus Elsemaria described from the rich Coniacian to Santonian flora of Hokkaido, Japan, exemplifies a permineralized capsular fruit of possible dilleniid affinity (NISHIDA, contribution 6). For age and sequence of stratigraphic names used in the palaeobotanical contributions readers may consult Fig. 4 in contribution 5 (DRINNAN & aI.). The neobotanical papers deal with floral development and biological aspects of previously poorly known magnoliids: Schisandra of Schisandraceae (TUCKER & BOURLAND, contribution 7), Barclaya of Nymphaeaceae (WILLIAMSON & SCHNEIDER, contribution 8), Cera to" phyllum of Ceratophyllaceae (ENDRESS, contribution 9, not read at the Symposium). Among ranunculids petal evolution is traced in Ranunculaceae (KOSUGE, contribu tion 10). Potential relationships of magnoliid flowers concerning outstanding features to those of other groups are discussed for monocots (ERBAR & LEINS, contribution 11) and for dilleniids and caryophyllids (LEINS & ERBAR, contribu tion 12). References BRENNER, G. J., 1963: The spores and pollen of the Potomac Group of Maryland. - Bull. Maryland Dept. GeoI. Mines Water Resources 27: 1-215. - 1976: Middle Cretaceous floral provinces and early migration of angiosperms. - In BECK, C. B., (Ed.): Origin and early evolution of angiosperms, pp. 23-47. - New York: Columbia University Press. CHASE, M. W., SOLTIS, D. E., OLMSTEAD, R. G., MORGAN, D., LES, D. H., MISHLER, B. D., DUVALL, M. R., PRICE, R A., HILLS, H. G., QIU, Y.-L., KRON, K. A., RETTIG, J. H., CONTI, E., PALMER, J. H., MANHART, J. R., SYTSMA, K. J., MICHAELS, H. J., KRESS, W. J., KAROL, K. G., CLARK, W. D., HEDREN, M., GAUT, B. S., JANSEN, R K., KIM, K.-J., WIMPEE, C. F., SMITH, J. F., FURNIER, G. R, STRAUSS, S. H., XIA1"G, Q.-Y., PLUNCKETT, G. M., SOLTIS, P. S., SWENSEN, S. M., WILLIAMS, S. E., GADEK, P. A., QUINN, C. J., EGUIARTE, L. E., GOLENBERG, E., LEARN, G. H., JR., GRAHAM, S. W., BARRETT, S. C. H., DAYANANDAN, S., ALBERT, V. A., 1993: Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid gene rbcL. - Ann. Missouri Bot. Gard. 80: 528-580. COEN, E. S., MEYEROWITZ, E. M., 1991: The war of the whorls: genetic interactions controlling flower development. - Nature 353: 31-37. CORNET, B., 1989a: The reproductive morphology and biology of Sanmiguelia lewisii, and its bearing on angiosperm evolution in the Late Triassic. - EvoI. Trends PI. 3: 25-51. Introduction - Major trends in the study of early flower evolution 5 1989b: Late Triassic angiosperm-like pollen from the Richmond Rift Basin of Virginia, U.S.A. - Palaeontograph. 213B: 37-87. CRANE, P. R., 1985: Phylogenetic relationships in seed plants. - Cladistics 1: 329-348. 1993: Time for the angiosperms. - Nature 366: 631-632. - LrDGARD, S., 1989: Angiosperm diversification and paleolatitudinal gradients in Cretace ous floristic diversity. - Science 246: 675-678. CRONQUIST, A., 1988: The evolution and classification of flowering plants. 2nd edn. - Bronx: New York Botanical Garden. DILCHER, D. L., CRANE, P. R., 1985: Archaeanthus: an early angiosperm from the Cenomanian of the western interior of North America. - Ann. Missouri Bot. Gard. 71: 351-383. DOYLE, J. A., 1969: Cretaceous angiosperm pollen of the Atlantic coastal plain and its evolutionary significance. - J. Arnold Arbor. 50: 1-35. - DONOGHUE, M. J., 1986: Seed plant phylogeny and the origin of angiosperms: an experimental cladistic approach. - Bot. Rev. 52: 321-431. 1993: Phylogenies and angiosperm diversification. - PaleobioI. 19: 141-167. - HICKEY, L. J., 1976: Pollen and leaves from the mid-Cretaceous Potomac Group and their bearing on early angiosperm evolution. - In BECK, C B., (Ed.): Origin and early evolution of angiosperms, pp. 139-206. - New York: Columbia University Press. ENDRESS, P. K., 1972: Zur vergleichenden Entwicklungsmorphologie, Embryologie und Systematik bei Laurales. - Bot. Jahrb. Syst. 92: 331-428. 1986: Reproductive structures and phylogenetic significance of extant primitive angio sperms. - PI. Syst. EvoI. 152: 1-28. 1990: Evolution of reproductive structures and functions in primitive angiosperms (Magnoliidae). - Mem. New York Bot. Gard. 55: 5-34. - HUFFORD, L. D., 1989: The diversity of stamen structures and dehiscence patterns among Magnoliidae. - Bot. J. Linn. Soc. 100: 45-85. FRIIS, E. M., CREPET, W. L., 1987: Time of appearance of floral features. - In FRIIS, E. M., CHALONER, W. G., CRANE, P. R., (Eds): The origins of angiosperms and their biological consequences, pp. 145-179. - Cambridge: Cambridge University Press. - ENDRESS, P. K., 1990: Origin and evolution of angiosperm flowers. - Adv. Bot. Res. 17: 99-162. - PEDERSEN, K. R., 1995: Angiosperm pollen in situ in Cretaceous reproductive organs. - In JANSONIUS, J., McGREGOR, D. C, (Eds): Palynology: principles and applications. - Amer. Assoc. Stratigraphic PalynoI. Foundation 1 (in press). - SKARBY, A., 1981: Structurally preserved angiosperm flowers from the Upper Cretaceous of southern Sweden. - Nature 291: 485-486. GOTTSBERGER, G., 1974: The structure and function of the primitive angiosperm flower - a discussion. - Acta Bot. Neerl. 23: 461-471. 1988: The reproductive biology of primitive angio~perms. - Taxon 37: 630-643. - SILBERBAUER-GOTTSBERGER, I., EHRENDORFER, F., 1980: Reproductive biology in the primitive relic angiosperm Drimys brasiliensis (Winteraceae). - PI. Syst. EvoI. 135: 11-39. HERENDEEN, P. S., 1991: Charcoalified angiosperm wood from the Cretaceous of eastern North America and Europe. - Rev. Palaeobot. PalynoI. 70: 225-239. HUGHES, N. F., 1976: Palaeobiology of angiosperm origins. - Cambridge: Cambridge University Press. JANSEN, R. K., PALMER, J. D., 1987: A chloroplast DNA inversion marks an ancient evolutionary split in the sunflower family (Asteraceae). - Proc. NatI. Acad. Sci. USA 84: 5818-5822. KATO, M., INOUE, T., 1994: Origin of insect pollination. - Nature 368: 195. KRUTZSCH, W., 1989: Paleogeography and historical phytogeography (paleochorology) in the Neophyticum. - PI. Syst. EvoI. 162: 5-61.

Description:
The recent discovery of a large number of excellently preserved fossil flowers, studied with new techniques, and the comparative study of the flowers of extant basal clades of the angiosperms revolutionized the conception of early flower evolution. This volume brings together contributions of 17 pal
See more

The list of books you might like

Most books are stored in the elastic cloud where traffic is expensive. For this reason, we have a limit on daily download.