The Botanical Review

, Volume 71, Issue 3, pp 354–414 | Cite as

Palynological characters and their phylogenetic signal in Rubiaceae

  • Steven Dessein
  • Helga Ochoterena
  • Petra De Block
  • Frederic Lens
  • Elmar Robbrecht
  • Peter Schols
  • Erik Smets
  • Stefan Vinckier
  • Suzy Huysmans


In the 1990s Rubiaceae became a hot spot for systematists, mainly due to the comprehensive treatment of the family by Robbrecht in 1988. Next to the exploration of macromolecular characters to infer the phylogeny, the palynology of Rubiaceae finally received the attention it deserves. This article aims to present a state-of-the-art analysis of the systematic palynology of the family. The range of varíation in pollen morphology is wide, and some of the pollen features are not known from other angiosperm taxa; e.g., a looplike or spiral pattern for the position of apertures in pantoaperturate grains. We compiled an online database at the generic level for the major pollen characters and orbicule presence in Rubiaceae. An overview of the variation is presented here and illustrated per character: dispersal unit, pollen size and shape, aperture number, position and type, sexine ornamentation, nexine pattern, and stratification of the sporoderm. The presence/absence and morphological variation of orbicules at the generic level is provided as well. The systematic usefulness of pollen morphology in Rubiaceae is discussed at the (sub)family, tribal, generic, and infraspecific levels, using up-to-date evolutionary hypotheses for the different lineages in the family. The problems and opportunities of coding pollen characters for cladistic analyses are also treated.


Botanical Review Pollen Type Pollen Morphology Pollen Character Pollen Size 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.

Literature Cited

  1. Adams, L. G., D. M. Bridson &E. Robbrecht. 1987. The identity ofLasianthus graciliflorus Bailey (Rubiaceae). Kew Bull. 42: 209–214.Google Scholar
  2. Aiello, A. 1979. A reexamination ofPortlandia (Rubiaceae) and associated taxa. J. Arnold Arbor. 60: 38–126.Google Scholar
  3. Ali, S. J. &E. Robbrecht. 1991. Remarks on the tropical Asian and Australian taxa included inDiplospora orTricalysia (Rubiaceae-Ixoroideae-Gardenieae). Blumea 35: 279–305.Google Scholar
  4. Anderson, W. R. 1972. A monograph of the genusCrusea (Rubiaceae). Mem. New York Bot. Gard. 22: 1–128.Google Scholar
  5. —. 1973. A morphological hypothesis for the origin of heterostyly in the Rubiaceae. Taxon 22: 537–542.Google Scholar
  6. Andersson, L. 1993. Pollen characteristics of the tribes Calycophylleae, Cinchoneae, and Hillieae (Rubiaceae). Nordic J. Bot. 13: 405–417.Google Scholar
  7. — &J. H. E. Rova. 1999. Therps16 intron and the phylogeny of the Rubioideae (Rubiaceae). Pl. Syst. Evol. 214: 161–186.Google Scholar
  8. Andreasen, K. &B. Bremen 2000. Combined phylogenetic analysis in the Rubiaceae-Ixoroideae: Morphology, nuclear and chloroplast DNA data. Amer. J. Bot. 87: 1731–1748.Google Scholar
  9. —,B. G. Baldwin &B. Bremen 1999. Phylogenetic utility of the nuclear rDNA ITS region in subfamily Ixoroideae (Rubiaceae): Comparisons with cpDNA rbcL sequence data. Pl. Syst. Evol. 217: 119–135.Google Scholar
  10. Andronova, N. N. 1984. The structure of the anther and pollen development in the Rubiaceae. Bot. Zhurn. 69: 43–54 (in Russian).Google Scholar
  11. Baker, H. G. 1956. Pollen dimorphism in the Rubiaceae. Evolution 10: 23–31.Google Scholar
  12. Bangoura, D. 1992. Beiträge zur Klärung der Gattungsabgrenzungsprobleme und Verwandtschaftsverhältnisse in der Tribus Pauridiantheae (Rubiaceae). Ph.D. diss., Univ. Wien, Vienna.Google Scholar
  13. Barrett, S. C. H. 1992. Heterostylous genetic polymorphisms: Model systems for evolutionary analysis. Pp. 1–29in S. C. H. Barrett (ed.), Evolution and function of heterostyly. Monographs on Theoretical and Applied Genetics, 15. Springer-Verlag, Berlin.Google Scholar
  14. Behnke, H. D. 1975. Electronenmikroskopische Untersuchungen zur Frage der verwandtschaftlichen Beziehungen zwischenTheligonum und Rubiaceae: Feinbau der Siebellement-Plastiden und Anmerkungen zur Struktur der Pollenexine. Plant. Syst. Evol. 123: 317–326.Google Scholar
  15. Bir Bahadur. 1963. Heterostylism inOldenlandia umbellata L. J. Genet. 58: 429–439.Google Scholar
  16. —. 1966. Heterostyly inOldenlandia scopulorum. Bull. J. Genet. 59: 267–272.Google Scholar
  17. —. 1968a. Pollen dimorphism in three heterostyled Rubiaceae. Rev. Palaeobot. Palynol. 7: 233–239.Google Scholar
  18. —. 1968b. Heterostyly in Rubiaceae: A review. Osmania Univ. J. Sci. [Golden Jubilee Vol.] 4: 207–238.Google Scholar
  19. —. 1970. Heterostyly inHedyotis nigricans (Lam.) Fosb. J. Genet. 60: 175–177.Google Scholar
  20. — &N. Rama Swamy. 1993. Pollen exine dimorphism in ten heterostylous species. J. Palynol. 29: 77–88.Google Scholar
  21. Borhidi, A. 1982. Studies in Rondeletieae (Rubiaceae), III. The generaRogiera andArachnothryx. Acta Bot. Acad. Sci. Hung. 28: 65–71.Google Scholar
  22. — &M. Fernández Zequeira. 1981a. Studies in Rondeletieae (Rubiaceae), I. A new genus:Roigella. Acta Bot. Acad. Sci. Hung. 27: 309–312.Google Scholar
  23. —. 1981b. Studies in Rondeletieae (Rubiaceae), II. A new genus:Suberanthus. Acta Bot. Acad. Sci. Hung. 27: 313–316.Google Scholar
  24. ——. 1983. Studies in Rondeletieae (Rubiaceae), V. Los límites del géneroSuberanthus. Acta Bot. Hung. 29: 29–34.Google Scholar
  25. —. 1983. Studies in Rondeletieae (Rubiaceae), IV. A new genus:Javorkaea. Acta Bot. Hung. 29: 13–27.Google Scholar
  26. —— &M. Moncada. 1980.Acunaeanthus, a new genus of Rubiaceae. Acta Bot. Acad. Sci. Hung. 26: 277–287.Google Scholar
  27. Bosser, J. &D. Lobreau-Callen. 1998.Landiopsis Capuron ex Bosser, genre nouveau de Rubiaceae de Madagascar. Adansonia, sér. 3, 20: 131–137.Google Scholar
  28. Bremekamp, C. E. B. 1952. The African species ofOldenlandia L. sensu Hiern et K. Schumann. Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect., 48: 1–297.Google Scholar
  29. —. 1963. On pollen dimorphism in heterostylous Psychotrieae, especially in the genusMapouria Aubl. Grana Palynol. 4: 53–63.Google Scholar
  30. —. 1966. Remarks on the position, the delimitation and the subdivision of the Rubiaceae. Acta Bot. Neerl. 15: 1–33.Google Scholar
  31. Bremer, B. &M. Thulin. 1998. Collapse of Isertieae, re-establishment of Mussaendeae, and a new genus of Sabiceeae (Rubiaceae): Phylogenetic relationships based on rbcL data. Pl. Syst. Evol. 211: 71–92.Google Scholar
  32. — &J. F. Manen. 2000. Phylogeny and classification of the subfamily Rubioideae (Rubiaceae). Pl. Syst. Evol. 225: 43–72.Google Scholar
  33. —,R. K. Jansen, B. Oxelman, M. Backlund, H. Lantz &K. J. Kim. 1999. More characters or more taxa for a robust phylogeny: Case study from the coffee family (Rubiaceae). Syst. Biol. 48: 413–435.Google Scholar
  34. Bridson, D. M. &E. Robbrecht. 1985a. Validation of the African genusHyperacanthus E. Mey. (Rubiaceae tribe Gardenieae). Kew Bull. 40: 273–286.Google Scholar
  35. ——. 1985b. Further notes on the tribe Pavetteae (Rubiaceae). Bull. Jard. Bot. Belg. 55: 83–115.Google Scholar
  36. —,P. Gasson &E. Robbrecht. 1980.Phellocalyx, a new tropical African genus in the Rubiaceae (Gardenieae). Kew Bull. 35: 315–321.Google Scholar
  37. Buchner, R. &C. Puff. 1993. The genus complexDanais-Schismatoclada-Payera (Rubiaceae): Character states, generic delimitation and taxonomic position. Bull. Mus. Natl. Hist. Nat., B, Adansonia 15: 23–74.Google Scholar
  38. Cabral, E. L. &N. M. Bacigalupo. 2001.Scandentia, nuevo género de Rubiaceae-Spermacoceae. Darwiniana 39: 29–41.Google Scholar
  39. Chaw, S. M. &S. P. Darwin. 1992. A systematic study of the paleotropical genusAntirhea (Rubiaceae: Guettardeae). Tulane Stud. Zool. Bot. 28: 25–118.Google Scholar
  40. Chennaveeraiah, M. S. &P. M. Shivakumar. 1983. Pollen bud formation and its role inOphiorrhiza spp. Ann. Bot. 51: 449–452.Google Scholar
  41. Chinnappa, C. C. &B. G. Warner. 1981. Pollen morphology in the genusCoffea (Rubiaceae) and its taxonomic significance. Bot. J. Linn. Soc. 83: 221–236.Google Scholar
  42. —. 1982. Pollen morphology in the genusCoffea (Rubiaceae), II. Pollen polymorphism. Grana 21: 29–37.Google Scholar
  43. Clément, C. &J.-C. Audran. 1993a. Orbicule wall surface characteristics in Lilium (Liliaceae). Grana 32: 348–353.Google Scholar
  44. —. 1993b. Electron microscope evidence for a membrane around the core of the Ubisch body in Lilium (Liliaceae). Grana 32: 311–314.Google Scholar
  45. Cortés-B., R. & S. Huysmans. Submitted. A survey of pollen and orbicules morphology ofRetiniphyllum (Rubiaceae-Ixorideae). Rev. Palaeobot. Palynol.Google Scholar
  46. Darwin, S. P. 1976. The genusLindenia (Rubiaceae). J. Arnold. Arbor. 57: 426–449.Google Scholar
  47. —. 1977. The genusMastixiodendron (Rubiaceae). J. Arnold. Arbor. 58: 349–381.Google Scholar
  48. —. 1979. A synopsis of the indigenous genera of Pacific Rubiaceae. Allertonia 2: 1–44.Google Scholar
  49. —. 1980a. Notes onAirosperma (Rubiaceae), with a new species from Fiji. J. Arnold. Arbor. 61: 95–105.Google Scholar
  50. —. 1980b.Habroneuron Standley, a little-known genus of Mexican Rubiaceae. Brittonia 32: 343–347.Google Scholar
  51. —. 1993. A revision ofTimonius subgenusTimonius (Rubiaceae: Guettardeae). Allertonia 7: 1–39.Google Scholar
  52. —. 1994. Systematics ofTimonius subgenusAbbottia (Rubiaceae: Guettardeae). Syst. Bot. Monogr. 42: 1–86.Google Scholar
  53. De Block, P. 1997. Biosystematic studies in the tribe Pavetteae (Rubiaceae-Ixoroideae). Ph.D. diss., Univ. Instelling Antwerpen, Antwerp.Google Scholar
  54. —. 1998. The African species ofIxora (Rubiaceae: Pavetteae). Opera Bot. Belg. 9: 1–218.Google Scholar
  55. —. 2003.Robbrechtia, a new Rubiaceae genus from Madagascar. Syst. Bot. 28: 145–156.Google Scholar
  56. — &E. Robbrecht. 1998. Pollen morphology of the Pavetteae (Rubiaceae, Ixoroideae) and its taxonomic significance. Grana 37: 260–275.Google Scholar
  57. —,J. Degreef &E. Robbrecht. 2001 [2002]. Reinstatement of the Afro-Malagasy genusCoptosperma (Rubiaceae, Ixoroideae, Pavetteae). Syst. Geogr. Pl. 71: 455–92.Google Scholar
  58. Delprete, P. G. 1999a. Rondeletieae (Rubiaceae). Part I. Fl. Neotrop. Monogr. 77: 1–226.Google Scholar
  59. —. 1999b.Riodocea (Rubiaceae, Gardenieae), a new genus from the Brazilian Atlantic forest. Brittonia 51: 15–23.Google Scholar
  60. Dessein, S. 2003. Systematic studies in the Spermacoceae (Rubiaceae). Ph.D. diss., Catholic Univ. of Leuven.Google Scholar
  61. —,A. Scheltens, S. Huysmans, E. Robbrecht &E. Smets. 2000. Pollen morphological survey ofPentas (Rubiaceae-Rubioideae) and its closest allies. Rev. Palaeobot. Palynol. 112: 189–205.PubMedGoogle Scholar
  62. —,S. Jansen, S. Huysmans, E. Robbrecht &E. Smets. 2001a. A morphological and anatomical survey ofVirectaria (African Rubiaceae), with a discussion of its taxonomic position. Bot. J. Linn. Soc. 137: 1–29.Google Scholar
  63. —,L. Andersson, E. Robbrecht &E. Smets. 2001b.Hekistocarpa (Rubiaceae): A member of an emended tribe Virectarieae. Pl. Syst. Evol. 229: 59–78.Google Scholar
  64. —,S. Huysmans, E. Robbrecht &E. Smets. 2002a. Pollen of AfricanSpermacoce species (Rubiaceae). Grana 41: 69–89.Google Scholar
  65. —,S. Jansen, E. Robbrecht &E. Smets. 2002b [2004]. A new species ofSpermacoce (Rubiaceae) from the Manika high plateau (Katanga; R. D. Congo). Nordic J. Bot. 22: 513–523.Google Scholar
  66. —,S. Ntore, E. Robbrecht &E. Smets. 2003. Pollen and seeds reveal thatSpermacoce thymoidea s. lat. (African Rubiaceae, Spermacoceae) represents three endemic or disjunct species from the Zambezian high plateaus. Syst. Bot. 28: 130–144.Google Scholar
  67. —,L. Andersson, K. Geuten, E. Smets &E. Robbrecht. 2005a.Gomphocalyx andPhylohydrax (Rubiaceae): Sister taxa excluded from the Spermacoceae s.s., featuring a remarkable case of convergent evolution. Taxon 54: 91–107.Google Scholar
  68. —,R. Harwood, S. Smets &E. Robbrecht. 2005b. Pollen of theSpermacoce species from the Northern Territory of Australia: Morphology and taxonomic significance. Austral. Syst. Bot. 18: 367–382.Google Scholar
  69. D’hondt, C. 2002. Pollen— en orbiculemorfologie van Hillieae (Rubiaceae). Ph.D. diss., Catholic Univ. of Leuven.Google Scholar
  70. —,P. Schols, S. Huysmans &E. Smets. 2004. Systematic relevance of pollen and orbicule characters in the tribe Hillieae (Rubiaceae). Bot. J. Linn. Soc. 146: 303–321.Google Scholar
  71. Di Maio, F. R. 1996 [1998]. Revisão taxonomica de generoHindsia Bentham (Rubiaceae, Hedyotideae). Arch. Jar. Bot. Rio de Janeiro 34: 51–92.Google Scholar
  72. El-Ghazaly, G &S. Huysmans. 2001. Re-evaluation of a neglected layer in pollen wall development with comments on its evolution. Grana 40: 3–16.Google Scholar
  73. — &E. Smets. 2001. Pollen wall development ofRondeletia odorata (Rubiaceae). Amer. J. Bot. 88: 14–30.Google Scholar
  74. Erdtman, G. 1971. Pollen morphology and plant taxonomy—Angiosperms. An Introduction to Palynology, I. Hafner Publishing, New York.Google Scholar
  75. Es, K. 1999.Geophila (Psychotrieae, Rubiaceae): Revisie van de neotropische taxa. Licentiate thesis, Catholic Univ. of Leuven.Google Scholar
  76. Farooq, M. &M. Inamuddin. 1969. The embryology ofOldenlandia nudicaulis Roth. J. Indian Bot. Soc. 48: 166–173.Google Scholar
  77. Farris, S. S. 1990. Phenetics in camouflage. Cladistics 6: 91–100.Google Scholar
  78. Groeninckx, I. 2005. Zoektocht naar de taxonomische positie vanMitrasacmopsis (Rubiaceae) op basis van moleculaire en morfologische data. Licentiate thesis, Catholic Univ. of Leuven.Google Scholar
  79. Gustafsson, C. G. R. 1998. The neotropicalRosenbergiodendron (Rubiaceae, Gardenieae). Brittonia 50: 452–466.Google Scholar
  80. — &C. Persson. 2002. Phylogenetic relationships among species of the neotropical genusRandia (Rubiaceae, Gardenieae) inferred from molecular and morphological data. Taxon 51: 661–674.Google Scholar
  81. Hansson, T. &G El-Ghazaly. 2000. Development and cytochemistry of pollen and tapetum inMitriostigma axillare (Rubiaceae). Grana 39: 65–89.Google Scholar
  82. Havard, A. &B. Verdcourt. 1987. A pollen survey ofTapiphyllum (Rubiaceae-Vanguerieae). Kew Bull. 42: 605–609.Google Scholar
  83. Hesse, M. 1986. Orbicules and the ektexine are homologous sporopollenin concretions in Spermatophyta. Pl. Syst. Evol. 153: 37–48.Google Scholar
  84. Hooker, J. D. 1873. Ordo LXXXIV. Rubiaceae. Pp. 7–151in G. Bentham & J. D. Hooker (eds.), Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata defirmata. London.Google Scholar
  85. Huang, T. C. 1972. Pollen flora of Taiwan, VI. National Taiwan Univ., Botany Dept. Press, Taipei.Google Scholar
  86. Huxley, C. R. &M. H. P. Jebb. 1993. The tuberous epiphytes of the Rubiaceae, 5. A revision ofMyrmecodia. Blumea 37: 271–334.Google Scholar
  87. Huysmans, S. 1993. De pollenmorfologie van de Coptosapelteae (Rubiaceae-Cinchonoideae). Licentiate thesis, Catholic Univ. of Leuven.Google Scholar
  88. -. 1998. Palynology of the Cinchonoideae (Rubiaceae): Morphology and development of pollen and orbicules. Ph.D. diss., Catholic Univ. of Leuven.Google Scholar
  89. —,E. Robbrecht &E. Smets. 1994. Are the generaHallea andMitragyna (Rubiaceae-Coptosapelteae) pollen morphologically distinct? Blumea 39: 321–340.Google Scholar
  90. —,G. El-Ghazaly, S. Nilsson &E. Smets. 1997. Systematic value of tapetal orbicules: A preliminary survey of the Cinchonoideae (Rubiaceae). Canad. J. Bot. 75: 815–826.Google Scholar
  91. —,E. Robbrecht &E. Smets. 1998a. A collapsed tribe revisited: Pollen morphology of the Isertieae (Cinchonoideae-Rubiaceae). Rev. Palaeobot. Palynol. 104: 85–113.Google Scholar
  92. —,G. El-Ghazaly &E. Smets. 1998b. Orbicules in Angiosperms: Morphology, function, distribution, and relation with tapetum types. Bot. Rev. (Lancaster) 64: 240–272.Google Scholar
  93. —,E. Robbrecht, P. Delprete &E. Smets. 1999. Pollen morphological support for the Catesbaeeae-Chiococceae-Exostema-complex (Rubiaceae). Grana 38: 325–338.Google Scholar
  94. —,G El-Ghazaly &E. Smets. 2000. Orbicules: Still a well hidden secret of the anther. Pp. 201–212in B. Nordenstam, G El-Ghazaly & M. Kassas (eds.), Plant systematics for the 21st century. Wenner-Gren International Series, 77. Portland Press, London.Google Scholar
  95. —,S. Dessein, E. Smets &E. Robbrecht. 2003. Pollen morphology of NW European representatives confirms monophyly of Rubieae (Rubiaceae). Rev. Palaeobot. Palynol. 127: 219–240.Google Scholar
  96. Igersheim, A. 1989. Beiträge zur Klärung der Gattungsabgrenzung-probleme innerhalb der Rubiaceae — Vanguerieae. Ph.D. diss., Botanical Institute, Vienna.Google Scholar
  97. —. 1991. Palynological investigations ofPaederia L. (Rubiaceae-Paederieae). Pp. 135–149in C. Puff (ed.), The genusPaederia L. (Rubiaceae-Paederieae): A multidisciplinary study. Opera Bot. Belg., 3. National Botanic Garden of Belgium, Meise.Google Scholar
  98. —. 1993a. The character states of the Caribbean monotypic endemicStrumpfia (Rubiaceae). Nordic J. Bot. 13: 545–559.Google Scholar
  99. —. 1993b. The palynology of the genusRondeletia L. (Rubiaceae-Cinchonoideae-Rondeletieae). Grana 32: 321–326.Google Scholar
  100. — &M. Weber. 1993 [1994]. “Pollen bud” formation inOphiorrhiza (Rubiaceae). Pp. 51–59in E. Robbrecht (ed.), Advances in Rubiaceae Macrosystematics. Opera Bot. Belg., 6. National Botanic Garden of Belgium, Meise.Google Scholar
  101. Jansen, S. 1994. Een houtanatomische en palynologische studie van Psychotrieae (Rubiaceae-Rubioideae). Licentiate thesis, Catholic Univ. of Leuven.Google Scholar
  102. —,E. Robbrecht &E. Smets. 1996a. The systematic value of endexine ornamentation in some Psychotrieae pollen (Rubiaceae-Rubioideae). Grana 35: 129–137.Google Scholar
  103. —,H. Beeckman &E. Smets. 1996b.Gaertnera andPagamea: Genera within the Psychotrieae or constituting the tribe Gaertnereae? A wood anatomical and palynological approach. Bot. Acta 109: 466–476.Google Scholar
  104. Johansson, J. T. 1987a. Pollen morphology of the tribe Morindeae (Rubiaceae). Grana 26: 134–150.Google Scholar
  105. —. 1987b. Revision of the genusPrismatomeris Thw. (Rubiaceae, Morindeae). Opera Bot. 94: 1–62.Google Scholar
  106. —. 1992. Pollen morphology inPsychotria (Rubiaceae, Rubioideae, Psychotrieae) and its taxonomic significance: A preliminary survey. Opera Bot. 115: 1–71.Google Scholar
  107. — &K. M. Wong. 1988. The identity ofPrismatomeris subsessilis King & Gamble (Rubiaceae, Rubioideae). Blumea 33: 351–356.Google Scholar
  108. Jung-Mendaçolli, S. L. 1984. Contribuição ao estudo palinológico das Rubiaceae. Ph.D. diss., Univ. of São Paulo.Google Scholar
  109. — &T. S. Melhem. 1994. Flora polinica da reserva do parque estadual das fontes do Ipiranga (São Paulo, Brasil). Hoehnea 21: 131–155.Google Scholar
  110. — &T. S. Melhem. 1995. Grãos de polen de especies heterostilicas de Rubiaceae (Pollen grains of heterostylous Rubiaceae). Revista Brasil. Bot. 18: 61–93.Google Scholar
  111. Keddam-Malplanche, M. 1980. Etude palynologique comparative des espèces lianescentes dans les genresSherbournia etPorterandia (Rubiacées-Gardéniées). Adansonia, sér. 2, 19: 429–434.Google Scholar
  112. -. 1985. Le pollen et les stomates des Gardeniées (Rubiacée) du Gabon. Morphologie et tendances évolutives. Mém. Mus. Nation. Hist. Nat. Nouv. Ser., Sér. B, 29. Paris.Google Scholar
  113. Kirkbride, J. H., Jr. 1979. Revision of the genusPsyllocarpus (Rubiaceae). Smithsonian Contr. Bot. 41: 1–32.Google Scholar
  114. Lecuona Neumann, R. M., I. La Serna Ramos, B. Mendes Pérez &W. Wildpret de la Torre. 1987. Contribución al estudio palinológico de la flora endémica macronesica. Pollen & Spores 29: 359–390.Google Scholar
  115. Lens, F., S. Jansen, S. Huysmans, E. Robbrecht &E. Smets. 2000. Pollen morphological variation in Vanguerieae (Ixoroideae-Rubiaceae). Grana 39: 1–13.Google Scholar
  116. Léonard, J. 1984. Contribution à la connaissance de la flore de l’Iran, VI. Le “complexeGaillionia A. Rich, ex DC.” (Rubiaceae). Bull. Jard. Bot. Belg. 54: 493–497.Google Scholar
  117. Leroy, J. F. 1974. Recherches sur les Rubiacees de Madagascar: Les genresMantalania etPseudomantalania (Gardeniées). Adansonia 14: 29–52.Google Scholar
  118. Lewis, W. H. 1965a. Cytopalynological study of African Hedyotideae (Rubiaceae). Ann. Missouri Bot. Gard. 52: 182–211.Google Scholar
  119. —. 1965b. Pollen morphology and evolution inHedyotis subgenusEdrisia (Rubiaceae). Amer. J. Bot. 52: 257–264.Google Scholar
  120. —. 1966. The Asian genusNeanotis nomen novum (Anotis) and allied taxa in the Americas (Rubiaceae). Ann. Missouri Bot. Gard. 53: 32–46.Google Scholar
  121. Lobreau-Callen, D. 1978. L’aperture composée des Rubiaceae. Annales Mines Belg. 2: 167–173.Google Scholar
  122. Lorence, D. H. 1986.Glossostipula (Rubiaceae), a new genus from Mexico and Guatemala. Candollea 41: 453–461.Google Scholar
  123. Luna, I. &H. Ochoterena. 2004. Phylogenetic relationships of the genera of Theaceae based on morphology. Cladistics 20: 223–270.Google Scholar
  124. Mabberley, D. J. 1997. The plant-book: A portable dictionary of the vascular plants. Ed. 2. Cambridge Univ. Press, Cambridge.Google Scholar
  125. Malplanche, M. 1971. Etude palynologique de trois genres de Rubiacées—Gardéniées d’Afrique. Adansonia, sér. 2, 11: 343–355.Google Scholar
  126. Mathew, P. M. &O. Philip. 1983. Studies in the pollen morphology of South Indian Rubiaceae. Vol. 10 of P. K. K. Nair (ed.), Advances in pollen-spore research. New Delhi: Today and Tomorrow’s Printers.Google Scholar
  127. —. 1987. Developmental and systematic significance of pollen bud formation inOphiorrhiza Linn. New Botanist 14: 47–54.Google Scholar
  128. McDowell, T. 1996.Syringantha coulteri (Hooker f.) T. McDowell, a new combination, and remarks on the relationships of the monotypic Mexican genusSyringantha Standley (Rubiaceae). Novon 6: 273–279.Google Scholar
  129. Motley T. J., K. J. Wurdack &P. G. Delprete. 2005. Molecular systematics of the Catesbaeeae-Chiococceae complex (Rubiaceae): Flower and fruit evolution and biogeographic implications. Amer. J. Bot. 92: 316–329.Google Scholar
  130. Muller, J. 1981. Fossil pollen records of extant angiosperms. Bot. Rev. (Lancaster) 47: 1–142.Google Scholar
  131. Naiki, A. & H. Nagamasu. 2003. Distyly and pollen dimorphism inDamnacanthus (Rubiaceae). J. Pl. Res. 105–113.Google Scholar
  132. Negron-Ortiz, V. 1996. Reproductive biology ofErnodea (Rubiaceae-Spermacoceae) in the Bahamas and Puerto Rico. Pp. 403–412in E. Robbrecht, C. Puff & E. Smets (eds.), Second Int. Rubiaceae Conf., Proceedings. Opera Bot. Belg., 7. National Botanic Garden Belgium, Meise.Google Scholar
  133. Nilsson, S. &J. Praglowski. 1992. Erdtman’s handbook of palynology. Ed. 2. Munksgaard, Copenhagen.Google Scholar
  134. Ntore, S. 2004. Contribution à la connaissance systématique du genre afrotropicalPauridiantha (Rubiaceae). Ph.D. diss., Catholic Univ. of Leuven.Google Scholar
  135. —,P. De Block, S. Huysmans, E. Robbrecht &S. Dessein. 2003. Two new species from Gabon show the need to reduceCommitheca to the synonymyof Pauridiantha (Rubiaceae, Pauridiantheae). Bot. J. Linn. Soc. 141: 105–117.Google Scholar
  136. Ochoterena, H. 2000. Systematics ofHintonia Bullock and thePortlandia complex (Rubiaceae). Ph.D. diss., Cornell Univ.Google Scholar
  137. Pacini, E. & G. G. Franchi. 1993. Role of the tapetum in pollen and spore dispersal. Pl. Syst. Evol., Suppl. 7: 1–11.Google Scholar
  138. —,G. G. Franchi &M. Hesse. 1985. The tapetum: Its form, function and possible phytogeny in embryophyta. Pl. Syst. Evol. 149: 155–185.Google Scholar
  139. Pailler, T. &J. D. Thompson. 1997. Distyly and variation in heteromorphic incompatibility inGaertnera vaginata (Rubiaceae) endemic to La Réunion Island. Amer. J. Bot. 84: 315–327.Google Scholar
  140. Persson, C. 1993. Pollen morphology of the Gardenieae-Gardeniinae (Rubiaceae). Nordic J. Bot. 13: 561–582.Google Scholar
  141. Philip, O. &P. M. Mathew. 1975. Cytology of exceptional development of the male gametophyte inOphiorrhiza mungos. Canad. J. Bot. 53: 2032–2037.Google Scholar
  142. Piesschaert, F. 2001. Carpology and pollen morphology of the Psychotrieae (Rubiaceae-Rubioideae): Towards a new tribal and generic delimitation. Ph.D. diss., Catholic Univ. of Leuven.Google Scholar
  143. —,S. Jansen, S. Huysmans, E. Smets &E. Robbrecht. 1999a.Chassalia petitiana (Rubiaceae-Psychotrieae), an overlooked epiphytic species hidden in the African canopy. Syst. Bot. 24: 315–322.Google Scholar
  144. —,E. Robbrecht, A. D. Poulsen &E. Smets. 1999b. Pyrene and pollen observations in the pantropical genusGeophila (Rubiaceae-Psychotrieae). Nordic J. Bot. 19: 93–100.Google Scholar
  145. — &E. Smets. 1999c.Chassalia subcordatifolia, a new combination in African Rubiaceae (Rubioideae, Psychotrieae). Syst. Geogr. Pl. 69: 189–194.Google Scholar
  146. —,L. Andersson, S. Jansen, S. Dessein, E. Robbrecht &E. Smets. 2000a. Searching for the taxonomic position of the African genusColletoecema (Rubiaceae): Morphology and anatomy compared to an rpsl6-intron analysis of the Rubioideae. Canad. J. Bot. 78: 288–304.Google Scholar
  147. —,S. Huysmans, I. Jaimes, E. Robbrecht &E. Smets. 2000b. Morphological evidence for an extended tribe Coccocypseleae (Rubiaceae-Rubioideae). Pl. Biol. 2: 536–546.Google Scholar
  148. —,S. Jansen, I. Jaimes, E. Robbrecht &E. Smets. 2001. Morphology, anatomy, and taxonomic position ofPagameopsis (Rubiaceae-Rubioideae). Brittonia 53: 490–504.Google Scholar
  149. Pire, S. M. 1996. Palynological study of American species ofBorreria (Rubiaceae-Spermacoceae). Pp. 416–423in E. Robbrecht, C. Puff & E. Smets (eds.), Second Int. Rubiaceae Conf., Proceedings. Opera Bot. Belg., 7. National Botanic Garden Belgium, Meise.Google Scholar
  150. —. 1997a. GéneroGalianthe subg.Ebelia (Rubiaceae: Spermacoceae): Estudio palinológico. Ann. Missouri Bot. Gard. 84: 878–887.Google Scholar
  151. -Pire, S. M.. 1997b. The pollen of Brazilian species ofRichardia L. (Rubiaceae-Spermacoceae). Rev. Univ. Guarulhos, Geosiências II Sp. Vol.: 184–191.Google Scholar
  152. — &E. L. Cabral. 1992. El valor del polen en la revalidación deGalianthe (Spermacoceae-Rubiaceae). Darwiniana 31: 1–10.Google Scholar
  153. Puff, C. 1986a. Abiosystematic study of the African and Madagascan Rubiaceae. Anthospermae. Springer-Verlag, ViennaGoogle Scholar
  154. —. 1986b.Phylohydrax (Rubiaceae-Spermacoceae)—A new genus to accommodate the African and Madagascan“Hydrophylax” species. Pl. Syst. Evol. 154, 343–366.Google Scholar
  155. —. 1988. Observations onCarphalea Juss. (Rubiaceae, Hedyotideae), with particular reference to the Madagascan taxa and its taxonomic position. Bull. Jard. Bot. Belg. 58: 271–323.Google Scholar
  156. —. 1989. The affinities and relationships of the Japanese endemicPseudopyxis (Rubiaceae: Paederieae). Pl. Spec. Biol. 4: 145–155.Google Scholar
  157. — &R. Buchner. 1998.Lecananthus andLeucocodon, two genera to be added to the tribe Schradereae (Rubiaceae). Blumea 43: 265–286.Google Scholar
  158. — &A. Igersheim. 1994a. The character states and taxonomic position ofMetabolos Bl. (syn.Allaeophania Thw.) (Rubiaceae). Bull. Jard. Bot. Belg. 63: 241–262.Google Scholar
  159. —. 1994b. The character states ofMussaendopsis Baill. (Rubiaceae-Coptosapelteae). Flora 189: 161–178.Google Scholar
  160. — &E. Robbrecht. 1988. The taxonomic position of the Australian endemicDurringtonia (Rubiaceae). Austral. Syst. Bot. 1: 191–197.Google Scholar
  161. —.. 1989. A survey of the Knoxieae (Rubiaceae-Antirheoideae). Bot. Jahrb. Syst. 110: 511–558.Google Scholar
  162. — &U. Rohrhofer. 1993 [1994]. The character states and taxonomic position of the monotypic mangrove genusScyphiphora (Rubiaceae). Pp. 143–172in E. Robbrecht (ed.), Advances in Rubiaceae macrosystematics. Opera Bot. Belg., 6. National Botanic Garden of Belgium, Meise.Google Scholar
  163. —,E. Robbrecht &V. Randrianasolo. 1984. Observations on the SE African-Madagscan genusAlberta and its allyNematostylis (Rubiaceae, Alberteae), with a survey of the species and a discussion of the taxonomic position. Bull. Jard. Bot. Belg. 54: 293–366.Google Scholar
  164. —,L. Andersson, U. Rohrhofer &A. Igersheim. 1993a. The tribe Schradereae (Rubiaceae) reexamined. Bot. Jahrb. Syst. 114: 449–479.Google Scholar
  165. —,A. Igersheim &U. Rohrhofer. 1993b.Pseudomussaenda andSchizomussaenda (Rubiaceae): Close allies ofMussaenda. Bull. Jard. Bot. Belg. 62: 35–68.Google Scholar
  166. —,R. Buchner &A. Igersheim. 1996.Dichilanthe, an unusual Asiatic Rubiaceae with “Lonicera flowers” and “dipterocarp fruits.” Nordic J. Bot. 16: 145–164.Google Scholar
  167. Punt, W., S. Blackmore, S. Nilsson & A. Le Thomas. 1994. Glossary of pollen and spore terminology. LPP Foundation, Utrecht. (Updated online version: /index.htm.)
  168. Puttock, C. F. 1992. Systematics of the Australian Gardenieae (Rubiaceae). Ph.D. diss., Univ. of New South Wales.Google Scholar
  169. Ramam, S. S. 1954. Gametogenesis and fertilization ofStephegyne parviflora Korth. Agra Univ. J. Res., Sci. 3: 243–348.Google Scholar
  170. Razafimandimbison, S. G. &B. Bremer. 2001 [2002]. Tribal delimitation of Naucleeae (Cinchonoideae, Rubiaceae): Inference from molecular and morphological data. Syst. Geogr. Pl. 71: 515–538.Google Scholar
  171. Reitsma, T. 1969. Size modifications of recent pollen grains under different treatments. Rev. Palaeobot. Palynol. 9: 175–202.Google Scholar
  172. Robbrecht, E. 1977. The tropical African genusHymenocoleus (Rubiaceae-Psychotrieae): Additions. Bull. Jard. Bot. Belg. 47: 3–29.Google Scholar
  173. —. 1978a.Sericanthe, a new African genus of Rubiaceae (Coffeeae). Bull. Jard. Bot. Belgique 48: 3–78.Google Scholar
  174. —. 1978b. Some observations inPreussiodora Keay (African Rubiaceae, Gardenieae). Bull. Soc. Roy. Bot. Belgique 111: 3–9.Google Scholar
  175. —. 1979. The African genusTricalysia A. Rich. (Rubiaceae-Coffeeae), 1. A revision of the species of subgenusEmpogona. Bull. Jard. Bot. Belg. 49: 239–360.Google Scholar
  176. -. 1980. Bijdragen tot de classificatie van de Ixoroideae en tot de revisie vanTricalysia s.l. (Rubiaceae). Ph.D. diss., Univ. Gent.Google Scholar
  177. —. 1981a. Studies in tropical African Rubiaceae, I.1. A new species ofTricalysia subgenusEmpogona from Gabon. 2. The position of“Tricalysia auriculata” and“Neorosea adamii.” 3. A new species ofBatopedina (Hedyotideae) from Shaba. 4. The distribution ofBatopedina andParapentas. Bull. Jard. Bot. Belg. 51: 165–189.Google Scholar
  178. —. 1981b. Studies in tropical African Rubiaceae, II. 5. A survey ofArgocoffeopsis. 6. A revision ofCalycosiphonia. Bull. Jard. Bot. Belg. 51: 359–378.Google Scholar
  179. —. 1982a. The African genusTricalysia A. Rich. (Rubiaceae-Coffeeae), 2.Ephedranthera, a new section of subgenusTricalysia. Bull. Jard. Bot. Belg. 52: 311–339.Google Scholar
  180. —. 1982b. Pollen morphology of the tribes Anthospermeae and Paederieae (Rubiaceae) in relation to taxonomy. Bull. Jard. Bot. Belg. 52: 349–366.Google Scholar
  181. —. 1983. The African genusTricalysia A. Rich. (Rubiaceae), 3.Probletostemon revived as a section for subgenusTricalysia. Bull. Jard. Bot. Belg. 53: 299–320.Google Scholar
  182. —. 1984. The delimitation and taxonomic position of the tropical African generaLeptactina andDictyandra (Rubiaceae). Pl. Syst. Evol. 145: 105–118.Google Scholar
  183. —. 1985. Further observations on the pollen morphology of the South African genusCarpacoce (Rubiaceae-Anthospermeae). Rev. Palaeobot. Palynol. 45: 361–371.Google Scholar
  184. —. 1987. The African genusTricalysia A. Rich. (Rubiaceae), 4. A revision of the species of sectionTricalysia. Bull. Jard. Bot. Belg. 57: 39–208.Google Scholar
  185. —. 1988a. Tropical woody Rubiaceae: Characteristic features and progressions: Contributions to a new subfamilial classification. Opera Bot. Belg., 1. National Botanic Garden of Belgium, Meise.Google Scholar
  186. —. 1988b. Studies in tropical African Rubiaceae, 13.Petitiocodon, a new genus to accommodateDidymosalpinx parviflora (Gardenieae-Diplosporinae). Bull. Jard. Bot. Belg. 58: 109–120.Google Scholar
  187. —. 1993 [1994]. Supplement to the 1988 outline of the classification of the Rubiaceae. Index to genera. Pp. 173–196in E. Robbrecht (ed.), Advances in Rubiaceae macrosystematics. Opera Bot. Belg., 6. National Botanic Garden of Belgium, Meise.Google Scholar
  188. — &D. M. Bridson. 1984. The taxonomic position of the East African genusCladoceras (Rubiaceae). Bull. Soc. Roy. Bot. Belgique 117: 247–251.Google Scholar
  189. — &C. Puff. 1981.Mericocalyx Bamps, synonymous withOtiophora Zucc. (Rubiaceae). Bull. Jard. Bot. Nat. Belg. 51: 143–151.Google Scholar
  190. — &C. Puff. 1986. A survey of the Gardenieae and related tribes. Bot. Jahrb. Syst. 108: 63–138.Google Scholar
  191. —— &A. Igersheim. 1991. Evidence for the close alliance betweenMitchella andDamnacanthus, with comments on the campylotropy in the Rubiaceae and the circumscription of the Morindeae. Blumea 35: 307–345.Google Scholar
  192. —,U. Rohrhofer &C. Puff. 1993 [1994]. A survey ofBertiera (Rubiaceae), including a discussion of its taxonomic position. Pp. 101–141in E. Robbrecht (ed.), Advances in Rubiaceae macrosystematics. Opera Bot. Belg., 6. National Botanic Garden of Belgium, Meise.Google Scholar
  193. —,S. Huysmans &E. Figueiredo. 1996. The generic status ofOxyanthus gossweileri (Rubiaceae) in Angola. S. African J. Bot. 62: 17–22.Google Scholar
  194. Rogers, G. K. 1984.Gleasonia, Henriquezia, andPlatycarpum (Rubiaceae). Fl. Neotrop. Monogr. 39: 1–135.Google Scholar
  195. Rova, J. 1999. The Condamineeae Rondeletieae Sipaneeae complex (Rubiaceae). Ph.D. diss., Univ. Göteborg.Google Scholar
  196. — &L. Andersson. 1995. A reevaluation of the tribes Hippotideae and Tammsieae (Rubiaceae). Nordic J. Bot. 15: 269–284.Google Scholar
  197. —,P. G Delprete, L. Andersson &V. A. Albert. 2002. A trnL-F cpDNA sequence study of the Condamineeae-Rondeletieae-Sipaneeae complex with implications on the phylogeny of the Rubiaceae. Am. J. Bot. 89: 145–159.Google Scholar
  198. Rutishauser, R., L. P. Ronse Decraene, E. Smets &I. Mcndoza-Heuer. 1998.Theligonum cynocrambe: Developmental morphology of a peculiar rubiaceous herb. Pl. Syst. Evol. 210: 1–24.Google Scholar
  199. Scheltens, A. 1998. Pollenmorfologische studie van de Afrikaanse Hedyotideae (Rubiaceae). Licentiate thesis, Catholic Univ. of Leuven.Google Scholar
  200. Schumann, K. 1891. Rubiaceae. Pp. 1–156in A. Engler & K. Prantl (eds.), Die natürlichen Pflanzenfamilien. Engelmann, Leipzig.Google Scholar
  201. Sotolongo Molina, L., M. Fernández Zequeira &P. Herrera Oliver. 2002. Pollen morphology of some CubanGuettarda species (Rubiaceae: Guettardeae). Grana 41: 142–148.Google Scholar
  202. Stoffelen, P., E. Robbrecht &E. Smets. 1997. Pollen morphology ofCoffea andPsilanthus (Rubiaceae-Coffeeae), mainly from Africa. Grana 36: 313–327.Google Scholar
  203. Strait, D. S., M. A. Moniz &P. T. Strait. 1996. Finite mixture coding: A new approach to coding con-tinuous characters. Syst. Biol. 45: 67–78.Google Scholar
  204. Swiderski, D. L., M. L. Zelditch &W. L. Fink. 1998. Why morphometrics is not special: Coding quantitative data for phylogenetic analyses. Syst. Biol. 47: 508–519.PubMedGoogle Scholar
  205. Tange, C. 1994.Neomussaenda (Rubiaceae), a new genus from Borneo. Nordic J. Bot. 14: 495–500.Google Scholar
  206. -Tange, C.. 1995. The identity ofSiderobombyx and a new species ofXanthophytum (Rubiaceae). Nordic J. Bot. 575–581.Google Scholar
  207. —. 1996a. Studies in SE Asiatic Rondeletieae, I: The West Malaysian endemic genusAleisanthia (Rubiaceae). Nordic J. Bot. 16: 571–578.Google Scholar
  208. —. 1996b. Studies in SE Asiatic Rondeletieae, II:Aleisanthiopsis (Rubiaceae), a new genus from Borneo. Nordic J. Bot. 16: 563–570.Google Scholar
  209. -Tange, C.. 1997. A revision of the genusMouretia (Rubiaceae). Nordic J. Bot. 123–132.Google Scholar
  210. -Tange, C.. 1998.Cyaneuron (Rubiaceae), a new genus from Borneo and Sulawesi. Nordic J. Bot. 147–158.Google Scholar
  211. Terrell, E. E., W. H. Lewis, H. Robinson &J. W. Nowicke. 1986. Phylogenetic implications of diverse seed types, chromosome numbers and pollen morphology inHoustonia (Rubiaceae). Amer. J. Bot., 73: 103–115.Google Scholar
  212. Thiele, K. 1993. The holy grail of the perfect character: The cladistic treatment of morphometric data. Cladistics 9: 275–304.Google Scholar
  213. Tilney, P. M. 1986. The taxonomic significance of anatomical and morphological characters in the southern African species ofCanthium Lam. (Rubiaceae). Ph.D. diss., Univ. of Pretoria.Google Scholar
  214. — &A. E. Van Wyk. 1997. Pollen morphology ofCanthium, Keetia andPsydrax (Rubiaceae: Vanguerieae) in southern Africa. Grana 36: 249–260.Google Scholar
  215. Tirvengadum, D. D. 1982.Ramosmania, a new monotypic genus of Mascarene Rubiaceae. Nordic J. Bot. 2: 323–327.Google Scholar
  216. —. 1984.Glionnetia, nouveau genre de Rubiacées (Rondeletiées) des Seychelles. Bull. Mus. Natl. Hist. Nat., B, Adansonia 6: 197–205.Google Scholar
  217. —. 1993.Larsenaikia, a new genus of the Rubiaceae from Australia. Nordic J. Bot. 13: 175–184.Google Scholar
  218. —. 1998. Novelties inRubiaceae from the limestone flora of southeast Asia. Biogeographica (The Hague) 74: 163–175.Google Scholar
  219. — &E. Robbrecht. 1985. Remarks on three Hypobathreae from Rodrigues, Seychelles and Sri Lanka. Nordic J. Bot. 5: 455–461.Google Scholar
  220. Tissot, C. &R.W. J. M. van der Ham. 1994. Seventh bibliographic index to the pollen morphology of Angiosperms. Trav. Sect. Sci. Techn. Inst. Franç. Pondichérie 27: 1–345.Google Scholar
  221. —,H. Chikhi &T.S. Nayar. 1994. Pollen of wet evergreen forests of the western Ghats, India. Trav. Sect. Sci. Techn. Inst. Franç. Pondichérie 35: 1–133.Google Scholar
  222. Van Campo, M. 1976. Patterns of pollen morphological variation within taxa. Pp. 125–137in I. K. Ferguson & J. Muller (eds.), The evolutionary significance of the exine. Linnean Soc. Symp. ser. 1. Acad. Press, London.Google Scholar
  223. —. 1978. La face interne de l’exine. Rev. Palaeobot. Palynol. 26: 301–311.Google Scholar
  224. Vanthournout, S. 2002.Geophila. Revisie van de paleotropische taxa. Licentiate thesis, Catholic Univ. of Leuven.Google Scholar
  225. Verdcourt, B. 1958. Remarks on the classification of the Rubiaceae. Bull. Jard. Bot. État. 28: 209–290.Google Scholar
  226. —. 1980. A conspectus ofPolysphaeria (Rubiaceae). Kew Bull. 35: 97–130.Google Scholar
  227. —. 1981. Notes on African Rubiaceae. Kew. Bull. 36: 493–557.Google Scholar
  228. —. 1983. Notes on Mascarene Rubiaceae. Kew Bull. 37: 521–574.Google Scholar
  229. —. 1987. Notes on African Rubiaceae-Vanguerieae. Kew Bull. 42: 123–199.Google Scholar
  230. Verellen, J. 2002. Palynologische studie en revisie vanCoptosapelta (Rubiaceae). Licentiate thesis, Catholic Univ. of Leuven.Google Scholar
  231. —,E. Smets &S. Huysmans. 2004. The remarkable genusCoptosapelta (Rubiaceae): Pollen and orbicule morphology and systematic implications. J. Pl. Res. 117: 57–68.Google Scholar
  232. -,S. Huysmans, S. G. Razafimandimbison & E. Smets. Submitted. Pollen morphology of the tribe Naucleeae s. lat. (Rubiaceae-Cinchonoideae) and its taxonomic and phylogenetic significance. Bot. J. Linn. Soc.Google Scholar
  233. Vinckier, S., S. Huysmans &E. Smets. 2000. Morphology and ultrastructure of orbicules in the subfamily Ixoroideae (Rubiaceae). Rev. Palaeobot. Palynol. 108: 151–174.PubMedGoogle Scholar
  234. Weber, M. &A. Igersheim. 1994. “Pollen buds” inOphiorrhiza (Rubiaceae) and their role in pollenkitt release. Bot. Acta 107: 257–262.Google Scholar
  235. Wodehouse, R. P. 1935. Pollen grains. McGraw-Hill, New York.Google Scholar
  236. Ying, T.-S., Y.-L. Zhang &D. E. Boufford. 1993. The endemic genera of seed plants of China. Science Press, Beijing.Google Scholar
  237. Young, B. A., J. Schulz-Schaeffer &T. W. Caroll. 1979. Anther and pollen development in malesterile intermediate wheatgrass plants derived from wheat × wheatgrass hybrids. Canad. J. of Bot. 57: 602–618.Google Scholar

Copyright information

© The New York Botanical Garden 2005

Authors and Affiliations

  • Steven Dessein
    • 1
    • 3
  • Helga Ochoterena
    • 2
  • Petra De Block
    • 3
  • Frederic Lens
    • 1
  • Elmar Robbrecht
    • 3
  • Peter Schols
    • 1
  • Erik Smets
    • 1
  • Stefan Vinckier
    • 1
  • Suzy Huysmans
    • 1
  1. 1.Laboratory of Plant Systematics, Institute of Botany and MicrobiologyCatholic University of LeuvenLeuvenBelgium
  2. 2.Instituto de BiologiaUniversidad Nacional Autónoma de MéxicoMexico CityMexico
  3. 3.National Botanic Garden of BelgiumDomein van BouchoutMeiseBelgium

Personalised recommendations