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Palynological diversity and major evolutionary trends in Cyperaceae

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Abstract

Pollen and orbicule morphology of 84 species, representing 52 genera from all tribes and subfamilies are investigated, in order to assess the systematic value of palynological data and to determine palynological evolutionary trends in Cyperaceae. A total of 90% of the species are examined for the first time with scanning electron microscopy. Pollen grains of Cyperaceae are oblate spheroidal to perprolate in shape, inaperturate to polyporate with opercula or pontopercula on pori or colpi. We distinguished seven different sexine ornamentation patterns. Orbicules occur in all species investigated. Pollen morphological variation within Cyperaceae is considerable and includes dispersal unit; number, location and degree of differentiation of apertural zones; and sexine ornamentation patterns. In subfamily Mapanioideae both tribes can be characterized by palynological synapomorphies. However, in subfamily Cyperoideae, the observed pattern of variation does not fit the most recent molecular phylogeny due to high levels of homoplasy and polymorphism in major pollen characters.

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Acknowledgments

This research was financially supported by the research council of K. U. Leuven (OT/05/35) and the Fund for Scientific Research—Flanders (Belgium) (F.·W. O.—G.0268.04). S. Vinckier was a postdoctoral fellow of F.·W. O. in the course of this study. A. M. Muasya acknowledges the Norwegian Council for Higher Education Programme for Development Research & Education (NUFU project 53/03) for funding and Flora Research Permit from CapeNature (AAA005-00054-0028). We thank Prof. P. Goetghebeur from the University of Ghent and Prof. J. Rammeloo, Director of the National Botanic Garden of Belgium (BR) for the supply of living material and herbarium specimens. We are grateful to Anja Vandeperre for technical assitance. Dr. Ochoterena and Steven Janssens are acknowledged for their assistance with the character optimization.

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Authors and Affiliations

  1. Laboratory of Plant Systematics, Institute of Botany and Microbiology, Kasteelpark Arenberg 31, P.O. Box 2437, 3001, Louvain, Belgium

    Anne Nagels, Suzy Huysmans, Alex Vrijdaghs & Erik Smets

  2. Botany Department, University of Cape Town, Private Bag X3, Rondebosch, 7701, South Africa

    A. Muthama Muasya

  3. National Herbarium of the Netherlands, Leiden University Branch, P.O. Box 9514, 2300 RA, Leiden, The Netherlands

    Erik Smets

  4. Center for Transgene Technology and Gene Therapy, Flanders Institute for Biotechnology, K.U. Leuven, Campus Gasthuisberg, Herestraat 49, 3000, Louvain, Belgium

    Stefan Vinckier

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  1. Anne Nagels
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  2. A. Muthama Muasya
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  3. Suzy Huysmans
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  4. Alex Vrijdaghs
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Corresponding author

Correspondence to Anne Nagels.

Appendices

Appendix 1: Species studied are listed alphabetically; fixed material (without asterisk), herbarium specimens are indicated with an asterisks and living material with double asterisks

  • Afrotrilepis pilosa (Ridl.) Gilly (*), Cameroon, P. Goetghebeur, 5182 (GENT).

  • Amphiscirpus nevadensis S. Watson (*), USA, J. Bouharmont, 19926, BR-S·P. 917869 (BR).

  • Arthrostylis aphylla R. Br., Australia, R.K. Harwood, RKH 1161.

  • Baumea rubiginosa Boeck., Australia, J.J. Bruhl and Hodges, JH 792.

  • Becquerelia cymosa Brogniart (*), Brazil, M. Luceno, 33B (GENT).

  • Bulbostylis hispidula (Vahl) R.W. Haines, Kenya, A.M. Muasya, AM 2466.

  • Capeobolus brevicaulis (C.B. Clarke), J. Browning, South Africa, A.M. Muasya, AM 2203.

  • Carex capitata L., Botanical Garden University Ghent, Belgium, P. Goetghebeur, PG 10466.

  • Carex elata Lowe, cultivated in the Botanical Garden of the Institute of Botany and Microbiology (K.U.Leuven) Belgium, A. Vrijdaghs, AV11.

  • Carex monostachya, A. Rich. (*), Senegal, C. Vanden Berghen, 8701 (BR)

  • Caustis flexuosa, R. Br. (*), USA, E.F. Constable, 26663 (BR).

  • Caustis recurvata Spreng. (*), Australia, P.K. Endress, 4381 (GENT).

  • Chorizandra cymbaria R. Br. (*), Australia, K.L. Wilson and K. Frank, 8954 (GENT).

  • Chorizandra enodis Nees, Australia, K.L. Wilson and K. Frank, 8922 (GENT).

  • Chrysitrix dodii C.B. Clarke, South Africa, A.M. Muasya, AM 2797.

  • Cladium mariscus (L.) Pohl, National Botanical Garden of Belgium, A. Vrijdaghs, AV06.

  • Coleochloa setifera (Ridl.) Gilly, Kenya, A.M. Muasya, AM 2464.

  • Costularia humbertii Bosser (*), Madagascar, J.S. Miller and P.P. Lowry, II4175 (GENT).

  • Courtoisina assimilis (Steud.) P. Maquet, Kenya, A.M. Muasya, AM 2124.

  • Cyperus alternifolius L. (**), Botanical Garden University Ghent Belgium, A. Vrijdaghs, 2001/1114.

  • Cyperus articulatus L., Kenya, A.M. Muasya, AM 2168.

  • Cyperus dubius Rottb., Kenya, A.M. Muasya, AM 2188.

  • Cyperus haspan L., Kenya, A.M. Muasya, AM 2135 (EA).

  • Cyperus hemisphaericus Boeckeler (*), Tanzania, E. Milne-Redhead and P. Taylor, 8053A (BR).

  • Cyperus laevigatus L., Botanical Garden University Ghent, Belgium, P. Goetghebeur, PG 10202.

  • Cyperus rotundus L., Kenya, A.M. Muasya, AM 2164.

  • Diplasia karatifolia L.C. Rich (*), Bolivia, R. Rueda, 921 (GENT).

  • Dulichium arundinaceum Britton, Botanical Garden University Ghent, Belgium, P. Goetghebeur, PG 9914.

  • Eleocharis acutangula (Roxb.) Schult., Kenya, A.M. Muasya, AM 2437.

  • Eleocharis palustris R. Br., Hortus Botanicus Lovaniensis Belgium, A. Vrijdaghs, AV07b.

  • Eriophorum latifolium Hoppe, Hortus Botanicus Lovaniensis Belgium, A. Vrijdaghs, AV04.

  • Everardia montana Ridley (*), Venezuela, P.E. Berry, O. Huber and J. Rosales, 4912 (GENT).

  • Exochogyne amazonica C.B. Clarke (*), Brasil, M. Aparecida da Silva, C. Proença, E. Cardoso and J.P. Paixão, 23.5.1994 (GENT).

  • Ficinia brevifolia Nees, South Africa, A.M. Muasya, AM 2205 (BOL, EA, K).

  • Ficinia capitellum Nees, South Africa, A.M. Muasya, AM 2206 (BOL, EA, K).

  • Ficinia dunensis Levyns, South Africa, A.M. Muasya, AM 2242.

  • Ficinia gracilis Schrad., Kenya, A.M. Muasya, AM 2571.

  • Ficinia minutiflora C.B. Clarke, South Africa, A.M. Muasya, AM 2257 (BOL, EA, K).

  • Ficinia polystachya Levyns, South Africa, A.M. Muasya, AM 2320.

  • Ficinia radiata Kunth, South Africa, A.M. Muasya, AM 2262 (BOL, EA, K).

  • Ficinia tristachya (Vahl) Nees, South Africa, A.M. Muasya, AM 2255.

  • Ficinia zeyheri Boeckeler, South Africa, A.M. Muasya, AM 2209 (BOL, EA, K).

  • Fimbristylis complanata (Retz.) Link, Kenya, A.M. Muasya, AM 2147.

  • Fimbristylis xyridis R. Br., Australia, R.K. Harwood, RKH 1162.

  • Fuirena abnormalis C.B. Clarke, Kenya, A.M. Muasya, AM 2192.

  • Fuirena leptostachya Oliver, Kenya, A.M. Muasya, AM 2136.

  • Gahnia lanigera (R. Br.) Benth. (*), Australia, B.J. Blaylock, 1227 (GENT).

  • Hellmuthia membranacea (Thunb.) R.W. Haines and Lye, South Africa, A.M. Muasya, AM 2792 (KUL).

  • Hypolytrum jenmanii C.B. Clarke subsp. jenmanii (*), Guyane, J.J. de Granville, F. Crozier, 13652 (GENT).

  • Isolepis antarctica (Willd.) Roem. & Schult., South Africa, A.M. Muasya, AM 2247 (BOL, EA, K).

  • Isolepis digitata Nees ex Schrad., South Africa, A.M. Muasya, AM 2258.

  • Isolepis prolifera (Rottb.) R. Br., South Africa, A.M. Muasya, AM 2265.

  • Isolepis setacea (L.) R. Br., Kenya, A.M. Muasya, AM 2547.

  • Kobresia myosuroides Fiori & Paoletti, Botanical Garden University Ghent, Belgium, P. Goetghebeur, PG 10009.

  • Kyllinga eximia C.B. Clarke, Kenya, A.M. Muasya, AM 2137.

  • Kyllinga flava C.B. Clarke, Kenya, Musili, MM 009.

  • Kyllinga polyphylla Thou. ex Link (**), Botanical Garden University Ghent, Belgium, A. Vrijdaghs, 2004/21768.

  • Kyllinga sp. Rottb., Kenya, A.M. Muasya, AM 2658.

  • Kyllingiella polyphylla, Kenya, A.M. Muasya, AM 2435.

  • Lagenocarpus rigidus (Kunth) Nees subsp. rigidus (*), French Guiana, D. Torida-Marbot, 329 (GENT).

  • Lepironia articulata (Retzius) Domin (*), Papua New Guinea, P. Goetghebeur and W. Vyverman, 6673 (GENT).

  • Lipocarpha nana (A. Rich.) Cherm., Kenya, A.M. Muasya, AM 2194.

  • Lipocarpha rehmannii (Ridl.) Goetgh., Kenya, A.M. Muasya, AM 3132.

  • Machaerina flexuosa (Böckeler) Kern (*), Madagascar, J.S. Miller and A. Randrianasdo, 4382 (GENT).

  • Mapania cutatrecasasii T. Koyama (*), Costa Rica, G. Herrera, 3282 (GENT).

  • Mapania linderi Hutchinson (*), Ivory Coast, C.C.H. Jongkind, 4435 (GENT).

  • Oxycarium cubense (Poepp. & Kunth) Palla, Kenya, Mwachala, M340.

  • Phylloscirpus acaulis (Philippi) Goetghebeur and D.A. Simpson subsp. pachycaulis (*), Equador, S. Laegaard, S. Dhooge and E. Jones, 21519 (GENT).

  • Pseudoschoenus inanis (Thunb.) Oteng- Yeboah, South Africa, A.M. Muasya, AM 3061.

  • Pycreus flavescens Beauv. ex Rchb. (**), Botanical Garden University Ghent, Belgium, A. Vrijdags, 2005/0401.

  • Pycreus mundtii Nees, Kenya, A.M. Muasya, AM 2156.

  • Pycreus sanguinolentus (Vahl) Nees, Kenya, A.M. Muasya, AM 2157.

  • Rhynchospora sp. Vahl, Australia, R.K. Harwood, RKH 1127.

  • Schoenoplectus senegalensis (Hochst. ex Steud.) Palla, Kenya, Malombe 40.

  • Schoenoxiphium lehmannii Kunth ex Steud., Kenya, Malombe, KG 96.

  • Schoenoxiphium sparteum C.B. Clarke, Kenya, A.M. Muasya, AM 2566.

  • Schoenus nigricans L., UK, K. De Wale, 1239 (GENT).

  • Scirpoides holoschoenus (L.) Sojak (**), Botanical Garden University Ghent, Belgium, A. Vrijdaghs, 2003/1536.

  • Scirpus sylvaticus L. (*), Botanical Garden University Ghent, Belgium, P. Goetghebeur, 5382 (GENT).

  • Scleria rugosa R. Br., Australia, R.K. Harwood, RKH 1143.

  • Scleria terrestris (L.) Fassett (**), Botanical Garden University Ghent, Belgium, A. Vrijdaghs, 21768.

  • Tetraria compar H.C. Taylor (*), South africa, H.C. Taylor, 9996 (GENT).

  • Trianoptiles solitaria (C.B. Clarke) Levyns, South Africa, A.M. Muasya, AM 3024

  • Trichophorum alpinum Pers. (*), USA, D. Collet, 670 (BR).

  • Uncinia rubra Colenso ex Boott, Botanical Garden University Ghent, Belgium, P. Goetghebeur, PG 9727.

Appendix 2: A detailed list of the characters and their states as defined for the optimization and the combined analysis. Characters 1, 2 and 9 are quantitative and continuous characters

  1. 1.

    Polar axis

    1. (0)

      0 (<18)

    2. (1)

      1 (<28)

    3. (2)

      2 (<41)

    4. (3)

      3 (<52)

    5. (4)

      4 (>52)

  2. 2.

    P/E

    1. (0)

      0 (<1)

    2. (1)

      1 (<1.4)

    3. (2)

      2 (<1.7)

    4. (3)

      3 (<2.3)

    5. (4)

      4 (>2.3)

  3. 3.

    Pollen shape

    1. (0)

      suboblate (0.75–0.88)

    2. (1)

      spheroidal (0.88–1.14)

    3. (2)

      subprolate (1.14–1.33)

    4. (3)

      prolate (1.33–2.00)

    5. (4)

      perprolate (>2.00)

  4. 4.

    Distal aperture

    1. (0)

      absent

    2. (1)

      1 ulcus

    3. (2)

      1 sulcus

  5. 5.

    Number lateral apertures

    1. (0)

      absent

    2. (1)

      3

    3. (2)

      4

    4. (3)

      5

    5. (4)

      6

    6. (5)

      >6

  6. 6.

    Shape lateral apertures

    1. (0)

      colpi

    2. (1)

      pori

    3. (2)

      variable

  7. 7.

    Distinctness of apertures

    1. (0)

      pontoperculum

    2. (1)

      operculum

  8. 8.

    Sexine ornamentation

    1. (0)

      microechinate

    2. (1)

      granulate–perforate

    3. (2)

      fossulate

    4. (3)

      negative microreticulate

    5. (4)

      fossulate–rugulate

    6. (5)

      psilate–rugulate

    7. (6)

      microreticulate

  9. 9.

    Orbicule diameter

    1. (0)

      0 (<0.3)

    2. (1)

      1 (<0.6)

    3. (2)

      2 (<0.9)

    4. (3)

      3 (<1.2)

    5. (4)

      4 (>1.2)

  10. 10.

    Orbicule shape

    1. (0)

      angular

    2. (1)

      irregular

    3. (2)

      spherical

    4. (3)

      doughnut-shaped

  11. 11.

    Orbicule ornamentation

    1. (0)

      smooth

    2. (1)

      microgranules

    3. (2)

      microechinae

    4. (3)

      striae

Appendix 3

Table 5.

Table 5 Data matrix with coded characters
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Nagels, A., Muasya, A.M., Huysmans, S. et al. Palynological diversity and major evolutionary trends in Cyperaceae. Plant Syst Evol 277, 117–142 (2009). https://doi.org/10.1007/s00606-008-0111-2

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