Takhtajan Armen. Flowering Plants
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Superorder PIPERANAE 61
morphology, and stipule development. Am. J. Bot. 88:
214–2150.
González F and DW Stevenson. 2000a. Gynostemium develop-
ment in Aristolochia (Aristolochiaceae). Bot. Jahrb. Syst.
122: 249–291.
González F and DW Stevenson. 2000b. Perianth development
and systematics of Aristolochia. Flora 195: 370–391.
González FA and Stevenson DW. 2002. A phylogenetic analy-
sis of the subfamily Aristolochioideae (Aristolochiaceae).
Rev. Acad. Colomb. Cienc. Exact. Fis. Nat. 26(98): 25–57.
González F, PJ Rudall, and CA Furness. 2001. Microsporogenesis
and systematics of Aristolochiaceae. Bot. J. Linn. Soc. 137:
221–242.
Gregory MP. 1956. A phyletic rearrangement in the
Aristolochiaceae. Am. J. Bot. 43: 110–122.
Guedes M. 1968. La feuille vegetative et perianthe dequelques
Aristolochia. Flora B 158: 167–179.
Hagerup O. 1961. The perianthium of Aristolochia elegans
Mast. Bull. Res. Council Israel IOD: 348–351.
Hegnauer R. 1960. Chemotaxonomische Betrachtungen: II.
Phytochemische Hinweise für die Stellung der Aristo-
lochiaceae im System der Dicotyledonen. Die Pharmazie 15:
634–642.
Holm T. 1926. Saururus cernuas L.: A morphological study.
Am. J. Sei., 5th ser., 12: 162–168.
Huber H. 1985. Samenmerkmale und Gliederung der
Aristolochiaceen. Bot. Jahrb. Syst. 107: 277–320.
Huber H. 1993. Aristolochiaceae. In: K Kubitzki, ed., The fami-
lies and genera of vascular plants, vol. 2, pp. 129–140.
Springer, Berlin/Heidelberg/New York.
Jacobsson-Stiasny E. 1918. Zur Embryologie der Aristo-
lochiaceae. Denkschr. Akad. Wiss. Wien, Math.-Naturw. Kl.,
95: 1–13.
Jaramillo MA and PS Manos. 2001. Phylogeny and patterns of
fl oral diversity in the genus Piper (Piperaceae). Am. J. Bot.
88: 706–716.
Jaramillo MA, PS Manos, and EA Zimmer. 2004. Phylogenetic rela-
tionships of the perianthless Piperales: Reconstructing the evolu-
tion of fl oral development. Int. J. Plant Sci. 165: 403–416.
Johnson DS. 1900. On the endosperm and embryo of Peperomia
pellucida. Bot. Gaz. 30: 1–11.
Johnson DS. 1902. On the development of certain Piperaceae
(Piper adunca, P medium, Heckeria umbellata). Bot. Gaz.
34: 321–340.
Johnson DS. 1914. Studies on the development of the Piperaceae:
II. The structure and seed development of Peperomia hispi-
dula. Am. J. Bot. 1: 323–329, 357–397.
Johri BM and SP Bhatnagar. 1955. A contribution to the mor-
phology and life history of Aristolochia. Phyto-morphology
5: 123–137.
Kamelina OP. 1997. An addition to the embryology of
Lactoridaceae and Fouquieriaceae. Bot. Zhurn. 82: 25–29 (in
Russian with English summary).
Kanta K. 1962. Morphology and embryology of Piper nigrum L.
Phytomorphology 12: 207–221.
Kelly LM. 1997a. Floral morphology, homology, and phyloge-
netic relationships in Aristolochiaceae. Am. J. Bot. 84(6):
207 (Abstract).
Kelly LM. 1997b. A cladistic analysis of Asarum
(Aristolochiaceae) and implications for the evolution of
herkogamy. Am. J. Bot. 84: 1752–1765.
Kelly LM. 1998. Phylogenetic relationships in Asarum
(Aristolochiaceae) based on morphology and ITS sequences.
Am. J. Bot. 85: 1454–1467.
Kelly LM and F González. 2003. Phylogenetic relationships in
Aristolochiaceae. Syst. Bot. 28: 236–249.
Kubitzki K. 1993. Lactoridaceae. In: K Kubitzki, ed. The
families and genera of vascular plants, vol. 2, pp. 359–361.
Springer, Berlin/Heidelberg/New York.
Lammers TG, TF Stuessy, and MO Silva. 1986. Systematic rela-
tionships of the Lactoridaceae: An endemic family of the
Juan Fernandez Islands, Chile. Plant Syst. Evol. 152: 3–4.
Lebot V and J Levesque. 1989. The origin and distribution of
Kava (Piper methysticum Forst.f., Piperaceae): A phy-
tochemical approach. Allertonia 5: 223–281.
Lei L-G and H-X Liang. 1998a. Floral development in dioecious
species and trends of fl oral evolution in Piper sensu lato.
Bot. J. Linn. Soc. 127: 225–237.
Lei L-G and H-X Liang. 1998b. Pollen morphology and its taxo-
nomic signifi cance of Piperaceae. Acta Bot. Yunn. 20:
427–433.
Lei L-G and H-X Liang. 1999. Variations in fl oral development
in Peperomia (Piperaceae) and their taxonomic implications.
Bot. J. Linn. Soc. 131: 423–431.
Lei L-G, Z-Y Wu, and H-X Liang. 2002. Embryology of Zippelia
begoniaefolia (Piperaceae) and it systematic relationships.
Bot. J. Linn. Soc. 140: 49–64.
Leinfellner W. 1953. Die hypopeltaten Brakteen von Peperomia.
Oesterr. Bot. Z. 100: 601–615.
Leins P and C Erbar. 1985. Ein Beitrag zur Blütenent-wicklung
der Aristolochiaceen, einer Vermittlergruppe zu den
Monokotylen. Bot. Jahrb. Syst. 107: 343–368.
Leins P and C Erbar. 1995. Das frühe Differenzierungsmuster in
den Blüten von Saruma henryi Oliv. (Aristolochiaceae). Bot.
Jahrb. Syst. 117: 365–376.
Leins P, C Erbar, and WA van Heel. 1988. Note on the fl oral
development of Thottea (Aristolochiaceae). Blumea 33:
357–370.
Liang H-X. 1991. Karyomorphology of Gymnotheca and phy-
logeny of four genera in Saururaceae. Acta Bot. Yunn. 13:
303–307.
Liang H-X. 1992. Study on the pollen morphology of
Saururaceae. Acta Bot. Yunn. 14: 401–404.
Liang H-X. 1994. On the systematic signifi cance of fl oral organo-
genesis in Saururaceae. Acta Phytotax. Sin. 32: 425–432.
Liang H-X. 1995. On the evolution and distribution in
Saururaceae. Acta Bot. Yunn. 17: 255–267.
Liang H-X and SC Tucker. 1989. Floral development in Gymno-
theca chinensis (Saururaceae). Am. J. Bot. 76: 806–819.
Liang H-X and SC Tucker. 1990. Comparative studies of the
fl oral vasculature in Saururaceae. Am. J. Bot. 77: 607–623.
Liang H-X and SC Tucker. 1995. Floral ontogeny of Zippelia
begoniaefolia and its familial affi nity: Saururaceae or
Piperaceae? Am. J. Bot. 82: 681–687.
Ling HX, KY Pan, and ZD Chen. 1996. Floral organogenesis in
Saururus chinensis (Saururaceae). Acta Phytotax. Sinica 34:
565–568.
Lorch JW. 1959. The perianth of Aristolochia: A new interpreta-
tion. Evolution 13: 415–416.
Majumdar GP and P Pal. 1961. Developmental studies: VI. The
morphology of the so-called stipule of Piper, etc. Proc. Nad.
Inst. Sci. India 27: 26–39.
62 Subclass I. MAGNOLIIDAE
Ma Jin-shuang. 1990. The geographical distribution and the system
of Aristolochiaceae. Acta Phytotax. Sinica 28(5): 345–355.
Mathew PJ and Mathew PM. 2001. Pollen morphology of some
members of Piperaceae and its bearing on the systematics
and phylogeny of the family. Rheedea 11(2): 65–78.
Mathew PJ, PM Mathew, and P Pushpangadan. 1999. Cytology
and its bearing on the systematics and phylogeny of the
Piperaceae. Cytologia (Japan). 64: 301–307.
Meeuse ADJ. 1971. Interpretative gynoecial morphology of the
Lactoridaceae and the Winteraceae: a re-assessment. Acta
Bot. Neerl. 20: 221–238.
Meeuse ADJ. 1972. Taxonomic affi nities between Piperales and
Polycarpicae and their implications in interpretative fl oral
morphology. Adv. Plant Morph. 1972: 3–27.
Meng SW, ZD Chen, DZ Li, and HX Liang. 2002. Phylogeny of
Saururaceae based on mitochondrial matR gene sequence
data. J. Plant Res. 115(1118): 71–76
Meng S-W, D-Z Li, H-Z Liang. 2001. The phylogeny of
Saururaceae based on 5.8S rDNA sequences. Acta Bot.
Yunn. 23: 309–312.
Meng SW, and H Liang. 1997. Comparative embryology on
Saururaceae. Acta Bot. Yunn. 19(1): 67–74.
Metcalfe CR. 1987. Lactoridaceae. In: CR Metcalfe, ed.
Anatomy of the dicotyledons, 2nd ed., vol. 3, pp. 147–151.
Claredon Press, Oxford.
Mi Qiu-wen and Yang Chun-shu. 1991. Pollen morphology of
Asarum in China. Acta Phytotax. Sinica 29(2): 164–171.
Miyoshi N and H Kato. 1982. Pollen morphology by means of
scanning electron microscope: 5. Angiospermae (Piperales,
Podostemonales). Jpn. J. Palynol. 28: 7–11.
Mohana Rao PR. 1989. Seed and fruit anatomy in Aristolochia
and Asarum with a discussion of the affi nities of
Aristolochiaceae. Swamy Bot. Club 6(3–4): 105–119.
Morawetz W. 1985. Beiträge zur Karyologie und Systematik
der Gattung Thottea (Aristolochiaceae). Bot. Jahrb. Syst.
107: 1–4.
Murty YS. 1958. Studies in the order Piperales: II. A contribu-
tion to the study of vascular anatomy of the fl ower of
Peperomia. J. Indian Bot. Soc. 37: 474–491.
Murty YS. 1959a. Studies in the order Piperales: III. A contribu-
tion to the study of fl oral morphology of some species of
Peperomia. J. Indian Bot. Soc. 38: 120–139.
Murty YS. 1959b. Studies in the order Piperales: V. A contribu-
tion to the study of fl oral morphology of some species of
Piper. VI. A contribution to the study of fl oral morphology
of Pothomorphe umbellata (L.) Miq. Proc. Indian Acad. Sci.
49B: 52–65, 82–85.
Murty YS. 1959c. Studies in the order Piperales: VII. A contri-
bution to the study of morphology of Saururus cernuus
L. J. Indian Bot. Soc. 38: 195–203.
Murty YS. 1960a. Studies in the order Piperales: 1. A contribu-
tion to the study of vegetative anatomy of some species of
Peperomia. Phytomorphology 10: 50–59.
Murty YS. 1960b. Studies in the order Piperales: VIII. A contri-
bution to the morphology of Houttuynia cordata Thunb.
Phytomorphology 10: 329–341.
Nair NC and KR Narayanan. 1962. Studies on the
Aristolochiaceae: I. Nodal and fl oral anatomy. Proc. Natl.
Inst. Sci. India 28B: 211–227.
Neinhuis C, KW Hilu, and T Borsch. 2000. Systematics
of Aristolochiaceae: Molecular evidence. Am. J. Bot.
87(Suppl. 6): 146.
Neinhuis C, S Wanke, KW Hilu, K Müller, and T Borsch. 2005.
Phylogeny of Aristolochiaceae based on parsimony, like-
lihood, and Bayesian analyses of trnL-trnF sequences. Plant
Syst. Evol. 250: 7–26.
Nickrent D, A Blarer, Y-L Qiu, DE Soltis, PS Soltis, and M
Zanis. 2002. Molecular data place Hydnoraceae with
Aristolochiaceae. Am. J. Bot. 89: 1809–1817.
Nikiticheva ZI. 1981. Embryological features of some Piperales.
Acta Soc. Bot. Polon. 50: 329–332.
Nikiticheva ZI. 1988. Piperaceae, Peperomiaceae. In:
A Takhtajan, ed. Comparative seed anatomy, vol. 2,
pp. 97–104. Nauka, Leningrad (in Russian).
Nikiticheva ZI, MS Yakovlev, and TA Plyushch. 1981. The
development of ovule, embryo sac, embryo, and endosperm
in some species of the genus Peperomia (Piperaceae). Bot.
Zhurn. 66: 513–523 (in Russian).
Nishida R, JD Weintraub, P Feeny and H Fukami. 1993.
Aristolochic acids from Thottea spp. (Aristolochiaceae) and
osmeterial secretions of Thottea-feeding troidine swallowtail
larvae (Papilionidae). J. Chem. Ecol. 19: 1587–1594,
Okada H. 1986. Karyomorphology and relationships in some
genera of Saururaceae and Piperaceae. Bot. Mag. Tokyo 99:
289–299.
Pant DD and R Banerji. 1965. Structure and ontogeny of sto-
mata in some Piperaceae. Bot. J. Linn. Soc. 59: 223–228.
Parmer VS, et al. 1998. Polyphenols and alkaloids from Piper
species. Phytochemistry 49: 1069–1078.
Pfl uge GM. 1956. Phyletic rearrangement in the Aristolochiaceae.
Am. J. Bot. 43: 110–122.
Plisko MA. 1988. Saururaceae. In: A Takhtajan, ed. Comparative
seed anatomy, vol. 2,pp. 93–96. Nauka, Leningrad (in
Russian).
Plyushch TA. 1982a. Ultrastructure of Peperomia blanda L.
(Piperaceae) embryo sac. Ukrain. Bot. Zhur. 39(4): 88–91 (in
Russian with English summary).
Plyushch TA. 1982b. Ultrastructure of Peperomia blanda
(Piperaceae) embryo sac in the process of fertilization.
Ukrain. Bot. Zhurn. 39 (6): 30–36 (in Russian with English
summary).
Pontieri V and TL Sage. 1997. Characterization of pollen/carpel
interaction following self and crosspollination in the paleo-
herb family: Saururaceae. Am. J. Bot. 84(Suppl. 6): 65.
Prakash N, JF Brown, and Y-H Wang. 1994. An embryological
study of kava, Piper metjysticum. Austral. J. Bot. 42:
231–237.
Quibell CH. 1941. Floral anatomy and morphology of Anemopsis
californica. Bot. Gaz. 102: 749–758.
Raju MVS. 1961. Morphology and anatomy of the Saururaceae:
1. Floral anatomy and embryology. Ann. Missouri Bot. Card.
48: 107–124.
Remizowa M, PJ Rudall, and D Sokoloff. 2005. Evolutionary
transitions among fl owers of perianthless Piperales: infer-
ences from infl orescence and fl ower development in the
anomalous species Peperomia fraseri (Piperaceae). Int. J.
Plant Sci. 166: 925–943.
Renuka C and K Swarupanandan. 1986. Morphology of the
fl ower in Thottea siliquosa and the existence of staminodes
in Aristolochiaceae. Blumea 31: 313–318.
Rohweder O and E Treu-Koene. 1971. Bau und morphologische
Bedeutung der Infl oreszenz von Houttuynia cordata Thunb.
(Saururaceae). Vierteljahrsschr. Naturf. Ges. Zürich 116:
195–212.
Superorder PIPERANAE 63
Sampson FB. 1995. Pollen morphology of Lactoridaceae: A
re-examination. Grana 34: 100–107.
Samuel R and W Morawetz. 1989. Chromosome evolution
within Piperaceae. Plant Syst. Evol. 166(1–2): 105–117.
Samuelson G. 1914. Über die Pollenentwicklung von Annona
und Aristolochia und ihre systematische Bedeutung. Svensk
Bot. Tidskr. 8: 181–189.
Sastrapradja S. 1968. On the morphology of the fl ower in
Peperomia (Piperaceae) species. Ann. Bogor. 4: 235–244.
Schmidt OC. 1935. Aristolochiaceae. In: A Engler und K Prantl,
eds. Die natürlichen Pfl anzenfamilien, 2nd ed., vol. 16b,
pp. 204–242. Engelmann. Prantl. Lepipzig..
Schmitz F. 1872. Die Blüten-Entwicklung der Piperaceen. Bot.
Abhand. Morphol. Physiol. 2(8): 1–74.
Semple KS. 1974. Pollination in Piperaceae. Ann. Missouri Bot.
Gard. 61: 868–871.
Skottsberg C. 1953. The vegetation of the Juan Fernandez
Islands. The Nat. Hist. of the Juan Fernandez and Easter Is.
2: 793–960.
Small JK. 1931. The wild pepper plants of continental United
States. J. New York Bot. Gard. 32: 210–223.
Solereder H. 1889. Beiträge zur vergleichenden Anatomie der
Aristolochiaceen. Engl. Bot. Jahrb. 10: 410–524.
Stuessy TF, DJ Crawford, GJ Anderson, and RJ Jenner. 1998.
Systematics, biogeography and conservation of Lactoridaceae.
Perspectives in Plant Evol. Syst. 1–2: 267–290.
Sugawara T. 1987. Chromosome number of Saruma henryi
Oliver (Aristolochiaceae). Bot Mag. Tokyo 100: 99–102.
Täckholm G and E Söderberg. 1918. Neue Beispiele der simul-
tanen und successiven Wandbbildung in den Pollen-
mutterzellen. Svensk Bot. Tidskr. 12: 189–201.
Tanaka H. 1979. Pollination in Saururus chinensis (Lour.) Baill.
J. Jpn. Bot. 54: 221–224.
Tebbs MC. 1993. Piperaceae. In: K Kubitzki, ed. The families
and genera of vascular plants, vol. 2, pp. 516–520. Springer,
Berlin/Heidelberg/New York.
Tobe H, TF Stuessy, PH Raven, and K Oginuma. 1993.
Embryology and karyomorphology of Lactoridaceae. Am. J.
Bot. 80: 933–946.
Trelease W. 1922. The peltate Peperomia of North America.
Bot. Gaz. 73: 133–146.
Trelease W and TG Yunker. 1950. The Piperaceae of northern
South America. Urbana.
Tucker SC. 1975. Floral development in Saururus cernuus
(Saururaceae): 1. Floral initiation and stamen development.
Am. J. Bot. 62: 993–1007.
Tucker SC. 1976. Floral development in Saururus cernuus
(Saururaceae): II. Carpel initiation and fl oral vasculature.
Am. J. Bot. 63: 289–301.
Tucker SC. 1979. Ontogeny of the infl orescence of Saururus
cernuus (Saururaceae). Am. J. Bot. 66: 227–236.
Tucker SC. 1980. Infl orescence and fl ower development in the
Piperaceae: 1. Peperomia. Am. J. Bot. 67: 686–702.
Tucker SC. 1981. Infl orescence and development in Houttuynia
cordata (Saururaceae). Am. J. Bot. 68: 1017–1032.
Tucker SC. 1982a. Infl orescence and fl ower development in the
Piperaceae: II. Floral ontogeny of Piper. Am. J. Bot. 69:
1389–1401.
Tucker SC. 1982b. Infl orescence and fl oral ontogeny in the
Pipera ceae: III. Floral ontogeny of Piper. Am. J. Bot. 69:
743–752.
Tucker SC. 1985. Initiation and development of infl orescence
and fl ower in Anemopsis californica (Saururaceae). Am. J.
Bot. 72: 20–31.
Tucker SC and AW Douglas. 1996. Floral structure, develop-
ment, and relationships of paleoherbs: Saruma, Cabomba,
Lactoris, and selected Piperales, pp. 141–175. In: DW
Taylor, LJ Hickey, eds. Flowering plant origin, evolution and
phylogeny. Chapman & Hall, New York.
Tucker SC, AW Douglas, and H-X Liang. 1993. Utility of onto-
genetic and conventional characters in determining phyloge-
netic relationships of Saururaceae and Piperaceae (Piperales).
Syst. Bot. 18 (4): 614–641.
Vovk AG and GA Komar. 1988. Aristolochiaceae. In: A Takhtajan,
ed. Comparative seed anatomy, vol. 2, pp. 105–111. Nauka,
Leningrad (in Russian).
Wagner R. 1907. Zur Kentnis des Saruma henryi Oliv. Oesterr.
Bot. Z. 57: 265–271.
Wanke S and C Neinhuis. 2006. Systematics of pipevines:
Combining morphological and fast-evolving molecular
characters to investigate the relationships within subfamily
Aristolochioideae (Aristolochiaceae). Int. J. Plant Sci. 167:
1215–1227.
Wanke S, M-S Samain, L Vanderschaeve, G Mathieu,
P Goetghebeur, and C Neinhuis. 2006. Phylogeny of the
genus Peperomia (Piperaceae) inferred from the trnK/matK
region (cpDNA). Plant Biol. 8: 93–102.
Wanke S, MA Jaramillo, T Borsch, M-S Samain, D Quandt, and
C Neinhuis. 2007. Evolution of Piperales – matK gene and
trnK intron sequence data reveal lineage specifi c resolution
contrast. Mol. Phyl. Evol. 42: 477–497.
Weberling F. 1970. Weitere Untersuchungen zur Morphologie
des Unterblattes bei den Dikotylen: V. Piperales. Beitr. Biol.
Pfl . 46: 403–434.
Weisse A. 1927. Zur Kenntnis von Blattstellung und
Blütensstand der Aristolochiaceen. Ber. Deutsch. Bot. Ges.
45: 229–244.
Wood CE. 1971. The Saururaceae in the southeastern United
States. J. Arnold Arbor. 52: 479–485.
Wu C-Y and K Kubitzki. 1993. Saururaceae. In: K Kubitzki,
ed. The families and genera of vascular plants, vol. 2,
pp. 586–588. Springer, Berlin/Heidelberg/New York.
Wyatt RI. 1955. An embryological study of four species of
Asarum. J. Elisha Mitchell Sei. Soc. 71: 64–82.
Yoshida O. 1957, 1959, 1960, 1961. Embryologische Studien
über die Ordnung Piperales. Parts 1–5. J. Coll. Arts Chiba
Univ. 2: 172–178, 1957; 295–303, 1959; 3: 56–60, 155–162,
1960; 311–316, 1961.
Yuncker TG. 1958. The Piperaceae: A family profi le. Brittonia
10: 1–7.
Zavada MS and JM Benson. 1987. First fossil evidence for the
primitive angiosperm family Lactoridaceae. Am. J. Bot. 74:
1590–1594.
Zavada MS and TN Taylor. 1986. Pollen morphology of
Lactoridaceae. Plant Syst. Evol. 154: 31–39.
Zhang S-S, S-B HO, and Y Wang. 1984. A study on the anatomy
of vegetative organs of the Gymnotheca Decne. (Saururaceae)
in relation to its systematic position. Acta Phytotax. Sinica
22: 49–52 (in Chinese).
Zhitkov VS. 1977. Forms of phyllotaxis in the genus Peperomia
Ruiz et Pav. And their morphogenesis. Bull. Mosc. Soc. Nat.,
Biol. ser. 82: 103–119 (in Russian with English summary).
64 Subclass I. MAGNOLIIDAE
Order 15. HYDNORALES
Terrestrial parasitic herbs living on the roots of various
trees and shrubs. Vegetative body consists of two kinds
of roots: the coarse rhizomelike pilot-roots, which tra-
verse the soil and appear to have a root cup, and endog-
enously emerging from the pilot-root small unbranched,
slender, lateral haustorial roots of limited growth, which
have a defi nite root cup. Vessels, when present, with
simple perforations. Parenchymatous tissues of the
pilot-root with scattered tanniniferous mucilage cells or
sometimes lysigenous mucilage canals or cavities con-
taining catechin. Sieve-element plastids of So-type, i.e.,
devoid of any prominent contents, diffi cult to recog-
nize, and of different sizes. Leaves absent. Flowers
solitary, arising endogenously from the pilot-roots,
rather large, usually bisexual (functionally unisexual in
Hydnora esculenta), actinomorphic, apetalous, mal-
odorous, cantharophilous. Sepals 3-4(5), thick and
fl eshy, with a coarse, cracked, brown exterior, valvate,
connate below. Stamens as many as sepals and opposite
to sepals in Hydnora sessile; anthers forming a lobed
ringlike synandrium on the calyx tube, the lobes oppo-
site the sepals. In Prosopanche the very short fi laments
arising from the calyx tube and the anthers connate to
form a dome or cap with a small central opening;
anthers elongate, extrorse, opening longitudinally.
Tapetum secretory. Microsporogenesis successive.
Pollen grains 2-celled, 1-colpate (Hydnora) or 2–3-
colpate or trichotomocolpate (Prosopanche), with
homogeneous ectexine, nearly psilate. Small, fl eshy
staminodia present between androecium and gynoe-
cium and alternate with the stamens in Prosopanche.
Gynoecium of 3(4-5) carpels; ovary inferior, unilocu-
lar, with numerous ovules embedded in much-branched
placentas suspended from the top of the ovary (in
Hydnora individual placentas are deeply intruded; in
Prosopanche radially arranged placental lamellae fi ll
the cavity of the ovary). Stigma is fl at with numerous
ridges in parallel series. Ovules numerous (ca. 35,000
in each ovary in Prosopanche americana), orthotro-
pous, unitegmic (single massive integument in Hydnora,
but scarcely differentiated from the placenta and recog-
nizable only in the micropylar area in Prosopanche),
tenuinucellate. Female gametophyte bisporic, Allium-
type (Prosopanche) or tetrasporic, Adoxa-type
(Hydnora). Endosperm cellular. Fruits rather massive,
with thick, leathery pericarp and fl eshy interior, edible
(the placental tissues contain much starch), in
Prosopanche bursting in circumscissile fashion when
ripe. Seeds very numerous, minute, with extremely
hard, exotestal seed coat and minute, undifferentiated
embryo surrounded by copious endosperm with poly-
saccharide food reserves and single layer of perisperm.
Closely related to the Aristolochiaceae, especially
to the Asaroideae, and have probably originated
directly from their immediate ancestors. The affi nity
with the Aristolochiaceae has been recently supported
by molecular data (Nickrent et al. 2002).
1. HYDNORACEAE
C. Agardh 1821. 2/18. Drier parts of Africa, Reunion,
Madagascar, and Saudi Arabia (Hydnora) as well as dry
regions of Central and South America (Prosopanche).
Hydnora, Prosopanche.
Bibliography
Bruch C. 1923. Coleopteros fertilizadores de Prosopanche bur-
meisteri De Bary. Physis 7: 82–88.
Burkart A. 1963. Nota sobre Prosopanche bonacinae Speg.
(Hydnoraceae) su area y parasitismo sobre algodon.
Darwiniana 12: 633–638.
Chodat R. 1915. Les especes du genre Prosopanche. Bull. Soc.
Bot. Geneve, ser. 2, 7: 65–66.
Chodat R. 1916. Hydnoraceae. Bull. Soc. Bot. Geneve, ser. 2, 8:
186–201.
Cocucci AE. 1965. Estudios en el genero Prosopanche
(Hydnoraceae): I. Kurtziana 2: 53–74.
Cocucci AE. 1975. Estudios en el genero Prosopanche
(Hydnoraceae): II. Kurtziana 8: 7–15.
Cocucci AE. 1976. Estudios en el genero Prosopanche
(Hydnoraceae): III. Kurtziana 9: 19–39.
Cocucci AE. 1983. New evidence from embryology in angio-
sperm classifi cation. Nord. J. Bot. 3: 67–73.
Cocucci AE. 1996. 1996. Prosopanche (Hydnoraceae): somatic
and reproductive structures, biology, systematics, phylogeny
and potentiality as a parasitic weed. In: MT Moreno, JC
Cubero, eds. Advances in parasitic plant research, pp. 178–
193. Junta de Andalucia, Sevilla.
Dastur RH. 1922. Notes on the development of the ovule,
embryo sac, and embryo of Hydnora ajricana.Trans. Roy
Soc. South Afr. 10: 27–31.
De Bary A. 1868. Prosopanche burmeuteri: Eine neue Hydnoreae
aus Südamerika. Abhandl. Naturf. Ges. Halle 10: 243–272.
Harms H. 1935. Hydnoraceae. In: A Engler und K Prantl, eds.
Die natürlichen Pfl anzenfamilien, vol. 16b, pp. 282–295.
Engelmann, Leipzig.
Kuijt J. 1969. The biology of parasitic fl owering plants.
University of California Press, Berkeley.
Superorder RAFFLESIANAE 65
Meijer W. 1993. Hydnoraceae. In: K Kubitzki, ed. The families
and genera of vascular plants, vol. 2, pp. 341–343. Springer,
Berlin/Heidelberg/New York.
Musselman LJ and JH Visser. 1989. Taxonomy and natural his-
tory of Hydnora (Hydnoraceae). Aliso12(2): 317–326.
Nickrent DL, A Blarer, Y-L Qiu, DE Soltis, PS Soltis, and
M Zanis. 2002. Molecular data place Hydnoraceae with
Aristolochiaceae. Am. J. Bot. 89: 1809–1817.
Schimper AFW. 1880. Die Vegetationsorgane von Prosopanche
burmeuteri. Abhandl. Naturf. Ges. Halle 15: 21–47.
Solms-Laubach HG. 1874. Über den Bau des Samens in den
Familien der Raffl esiaceae und Hydnoraceae. Bot. Z. 32 (I):
337–342, 353–358, 369–374, 385–389.
Takhtajan AL, NR Meyer, and VN Kosenko. 1979. Morphology
of pollen grains of the family Hydnoraceae in relation to its
systematic position. Bot. Zhurn. 64: 1774–1777 (in Russian
with English summary).
Terekhin ES. 1981. Hydnoraceae. In: MS Yakovlev, ed.
Comparative embryology of fl owering plants, pp. 95–96.
Nauka, Leningrad (in Russian).
Terekhin ES. 1988. Hydnoraceae. In: A Takhtajan, ed.
Comparative seed anatomy, vol. 2, pp. 112–113. Nauka,
Leningrad (in Russian).
Tieghem P van. 1897. Sur la structure de 1’ovule et de la graine
chez les Hydnoracees. Jour. de Bot. 11: 233–238.
Visser J. 1981. South African parasitic plants. Juta, Capetown.
Superorder RAFFLESIANAE
Order 16. MITRASTEMONALES
Plants without chlorophyll, endoparasitic in the roots
of other plants, often causing broomlike overgrowth of
the host-root. Vegetative body much-dissected and
largely fi lamentous, resembling a fungal mycelium,
permeating the tissues of the root of the host, but not
extending into the apical meristem. Leaves opposite,
scale-like but sometimes fl eshy and leaf waxes hum-
mocky, the uppermost leaves tending to be somewhat
cupped and accumulating nectar. Stomata absent.
Flowers of medium size, terminal, solitary, bisexual,
protandrous. Nectaries in the axils of the upper bracts.
Perianth much reduced, collar-shaped, connate below
to form a cup; the rim of the cap 4-lobed. Stamens con-
nate into a tube (androphore); the staminal tube, open
at the top by a small hole, circumscissaly separated for
the fl ower as it is pushed up by the glowing gynoe-
cium. The apical portion of the staminal tube is sterile.
Anthers numerous, extrorse, sessile in several series of
rings shortly below the stamen tube summit, 2-locular,
opening longitudinally. Tapetum secretory. Micro-
sporogenesis simultaneous. Pollen grains 2-celled,
3(4)-porate, ectexine reduced to tuberculae. Gynoecium
with 9–15 intrusive parietal placentae; style short,
stout, the stigma thick, depressed-conical. Ovary supe-
rior, 1-locular. Ovules numerous, anatropous, uniteg-
mic (but integument with two cells layers),
tenuinucellate, with a funicular obturator. Female
gametophyte of Polygonum-type. Endosperm cellular.
Fruits baccate or capsular and tardily opening by a
horizontal slit. Seeds very numerous, minute; exotestal
cells with U thickening; embryo undifferentiated,
4-celled, surrounded by endosperm; n = 20.
Usually including in the Raffl esiales, but differ
from them in many respects, especially in superior
ovary and cellular endosperm.
1. MITRASTEMONACEAE
Makino 1911. 1/2. Eastern Asia and Southeast Asia,
Malesia, and America from Mexico to Guatemala and
northwestern Colombia.
Mitrastemon.
Bibliography
Jochems SCJ. 1928. Die Verbreitung der Raffl esiaceengattung
Mitrastemon. Rec. Trav. Bot. Neerl. 25A: 203–207.
Makino T. 1911. Mitrastemon – Mitrastemonaceae –
Mitrastemonales. Bot. Mag. Tokyo 25: 251–257.
Matuda E. 1947. On the genus Mitrastemon. Bull. Torrey Bot.
Club 74: 133–141.
Meijer W and JF Veldkamp. 1993. A revision of Mitrastema
(Raffl esiaceae). Blumea 38: 221–229.
Mitra K, M Mondal. 1982. A note on the pollen morphology of
Mitrastemon yamamotoi (Makino) Makino (Raffl esiaceae).
Bangladesh J. Bot. 11(2): 179–181.
Nickrent DL, A Blarer, Q Yin-Long, R Vidal-Russell, and FE
Anderson. 2004. Phylogenetic inference in Raffl esiales: the
infl uence of rate heterogeneity and horizontal gene transfer.
BMC Evol. Biol. 4: 40 – http://www.biomedcentral.
com/14711–2148/4/40
Watanabe K. 1936. Morphologisch-biologische Studien liber die
Gattung Mitrastemon: IV. J. Jpn. Bot. 12: 848–858.
Watanabe K. 1937. Morphologisch-biologische Studien über die
Gattung Mitrastemon: VII. J. Jpn. Bot. 13: 154–162.
Order 17. RAFFLESIALES (CYTINALES)
Parasitic herbs, often fl eshy, living on the roots and
less often the stems of various trees and shrubs.
66 Subclass I. MAGNOLIIDAE
External roots and rhizomelike structures absent.
The vegetative body is much branched, and the
myceliumlike endophytic system is inside the tis-
sues of the host. A very reduced, vestigial vascular
system exists only in the fairly massive haustorial
system and in the fl ower-bearing axis. The conduct-
ing elements are short, broad, spirally thickened tra-
cheids, but the older elements are mostly devoid of
transverse walls. So-plastids diffi cult to recognize
and of varying sizes. Reduced scalelike leaves pres-
ent on the emergent fl owering shoot or around the
base of the solitary fl ower, usually verticillate,
sometimes opposite or even alternate. Stomata occur
very rarely and are abnormal, mostly absent. Flowers
from very small (Apodanthaceae) to very large or
even gigantic (Raffl esiaceae), solitary and sessile or
less frequently in short racemes or spikes, bisexual
or more often unisexual (monoecious or dioecious),
apetalous, actinomorphic, often malodorous. Sepals
4–5, sometimes up to 10 or more, in 1 or 2 whorls,
free or more often connate below, sometimes peta-
loid. Stamens fi ve to many (up to 100), connate by
their fi laments into a tube surrounding the stylar
column, or more often adnate to the column, from
which the anthers originate in one to several cycles;
anthers unilocular or bilocular, tetrasporangiate,
opening longitudinally or by transverse slits or by
apical pore. Tapetum secretory. Microsporogenesis
simultaneous. Pollen grains 2-celled, monocolpate,
3-colpate, or 1-polyporate or inaperturate; ectexine
from homogeneous to columellate. Nectary some-
times present, adnate to the base of the style and
base of the staminal column. Gynoecium of 4-10(-
14) carpels; ovary inferior or semi-inferior, 1-locu-
lar with more or less intruded parietal placentas or
irregularly multilocular with the ovules covering the
surfaces of the placental partitions; stigma on a short
style or sessile, distally expanded into an often large
disc, capitate or multilobed, more or less papillose.
Ovules very numerous, hemianatropous to anatro-
pous or orthotropous (Cytinaceae), mostly bitegmic,
usually with reduced short outer integument, tenui-
nucellate. Female gametophyte of Polygonum-type.
Endosperm nuclear. Fruits usually more or less
berry-like, indehiscent, dehiscent by horizontal slit,
or irregularly dehiscent. Seeds very numerous, tiny,
with an undifferentiated embryo surrounded by
1(-3)-layered endosperm; seed-coat hard, of a single
layer. Perisperm absent.
Related to the Aristolochiaceae. The relationship
with the Aristolochiaceae has been accepted by Robert
Brown (1821), Solms-Laubach (1901), Hallier (1905b,
1908, 1912), Endress (1988, 1994), Bouman and
Meijer (1994), and many others. According to Hallier,
the family Raffl esiaceae is closest to the section
Heterotropa of the genus Asarum and originated from
the Aristolochiaceae as a result of parasitic reduction.
Key to Families
1 Ovules orthotropous. Flowers of medium size, in
clusters or spike-like infl orescences, unisexual
(monoecious or dioecious). Stems stout with bract-
like leaves. Flowers with bracts and bracteoles, with
more or less horseshoe-shaped nectary glands
(Cytinus), Stamens 8–10, with united fi laments,
often with appendages above the anthers. Anthers
2-locular. Pollen grains free or in tetrads, 2–4-porate
or 1–5-colpate, rugate or polyporate. Nectary disc
present (Cytinus). Ovary inferior, with 6–8 parietal
placentas. Style 1 with punctuate-radiate stigma or
stigma disk-like (Bdallophyton) Ovules unitegmic,
but with reduced outer integument (Tereckin 1988),
micropyle endostomal. Contain pelargonidin
3-galactoside and petunidin 3-glucoside (Hegnauer
1997), n = 16. . . . . . . . . . . . . . . . . . 3. cytinaceae
1 Ovules hemianatropous to anatropous.
2 Pollen grains triporate. Ovules bitegmic, anatro-
pous, covering the whole of inner wall of the
ovary or on 4–5 wide placentas. Endoparasitic,
achlorophyllous, monoecious or dioecious herbs;
their vegetative endophyte, often compared to a
fungal mycelium, residents in the host; Apodanthes
is known to parasitize Flacourtiaceae Pilostyles
and Berlinianche para sitize a wide range of
Fabaceae. Flowers small, not more than 4–5 mm
in diameter, unisexual. Staminal structures later-
ally on apex of the central column. Stamens ca.
15, in 2–4 cycles. Anthers 1-locular, extrorse,
opening by transverse slits. Pollen grains 3-col-
pate. Nectary disc present. Female fl owers cone-
like and broader than the male ones, with annular
stigma below the apex of the column. Gynoecium
of 4–5 united carpels; carpels opposite inner pet-
als; Stigma hemispherical without separate stig-
matic lobes and is almost completely covered
with elongate unicellular papillae, each with a
rounded apex, which are heavily secretory
(Berlinianche – Blarer, Nickrent, and Endress
Superorder RAFFLESIANAE 67
2004). Ovary inferior or semi-inferior, 1-locular,
placentation parietal; ovules many, anatropous,
micropyle bistomal or none. Fruits baccate, testa
thin-walled, exotegmen massively lignifi ed;
endosperm present, embryo undifferentiated;
n = 16, 30–31 + (Pilostyles). 1. apodanthaceae
2 Pollen grains unisulcate or uniporate. Flowers
very large, solitary or in infl orescences, uni-
sexual or (Rhizanthes) bisexual. Staminal struc-
tures under the rim of the expanded apex of the
central column. Stamens in one cycle. Anthers
with two to many locules. Pollen grains 1-porate
or with reduced short colpi. Raffl esia and
Sapria have no nectarines, but Rhizanthes has a
nectary on the distal part of the perianth organs
(Bänziger and Hansen 2000). Female fl owers
with stigma on or under the rim of apical disc.
Ovary inferior. Ovules basically anatropous,
unitegmic. Seeds with basal chalazal append-
age. n = 11, 12.. . . . . . . . . . 2. rafflesiaceae
1. APODANTHACEAE
Tieghem ex Takhtajan 1987. 3/23–30. America from
southern California and southeastern USA to the Straits
of Magellan, East Africa, western Asia (Asia Minor,
Iraq, Iran), southwestern Australia.
Apodanthes, Pilostyles, Berlinianche.
Related to the Raffl esiaceae, but differ in pollen
grains and bitegmic ovules.
2. RAFFLESIACEAE
Dumortier 1829. 3/18–20. Southern China, Assam,
Bhutan, Thailand, and Indochina (Sapria himalayana)
and Malesia.
Raffl esia, Sapria, Rhizanthes.
3. CYTINACEAE
A. Richard 1824. 2/8. South Africa, Madagascar,
Mediterranean, Asia Minor, and western Caucasus
(Cytinus), Central America (Bdallophytori).
Bdallophyton. Cytinus
Related to the Raffl esiaceae, but differ in orthotro-
pous ovules.
Bibliography
Bänziger H. 1995. Ecological, morphological and taxonomic
studies on Thailand fi fth species of Raffl esiaceae: Rhizanthes
zippelii (Blume) Spach. Nat. Hist. Bull. Siam Soc. 43:
337–365.
Bänziger H and B Hansen. 2000. A new taxonomic revision of a
deceptive fl ower, Rhizanthes Dumortier (Raffl esiaceae). Nat.
Hist. Bull. Siam Soc. 48: 117–143.
Beaman RS, PJ Decker, and JH Beaman. 1988. Pollination in
Raffl esia (Raffl esiaceae). Am. J. Bot. 75: 1148–1162.
Blarer A, DL Nickrent, and PK Endress. 2004. Comparative fl oral
structure and systematics in Apodanthaceae (Raffl esiales).
Plant Syst. Evol. 245: 119–142.
Blarer A., DL Nickrent, H Bänziger, PK Endress, and Y-L Qui.
2000. Phylogenetic relationships among genera of the para-
sitic family Raffl esiaceae s.l. nased pm mic;ear ITS and SSU
rDNA, mitochondrial LSU and SSU rDNA, atp1, and matR
sequences. Am. J. Bot. 87(Suppl. 6): 503.
Blarer A, DL Nickrent, and PK Endress. 2004. Comparative fl o-
ral structure and systematics in Apodanthaceae (Raffl esiales).
Plant Syst. Evol. 245: 119–142.
Bouman F and W Meijer. 1986. Comparative seed morphology
in Raffl esiaceae. Acta Bot. Neerl. 35: 521–521.
Bouman F and W Meijer. 1994. Comparative structure of ovules
and seeds in Raffl esiaceae. Plant Syst. Evol. 193: 187–212.
Brown R. 1821. An account of a new genus of plants named
Raffi esia. Trans. Linn. Soc. London 13: 231–234.
Brown R. 1834. Description of the female fl ower and fruit of
Raffl esia arnoldi, with remarks on its affi nities and an illus-
tration of the structure of Hydnora africana. Trans. Linn.
Soc. London 19: 221–247.
Cartellieri E von. 1926. Das Absorptionssystem der Raffl esiaceae
Brugmansia. Bot. Arch. 14: 284–311.
Chodat R and C Bernard. 1902. Embryologie du Cytinus hypo-
cistis. Arch. Sci. Phys. et Nat. Geneve, 13: 1–6.
Davis CC, M Latvis, DL Nickrent, KJ Wurdack, and DA
Baum. 2007. Floral gigantism in Raffl esiaceae. Science
315: 1812.
Dell B, J Kuo, and A Brium. 1982. Anatomy of Pilostyles ham-
iltonii C. L. Gardner (Raffl esiaceae) in stems of Daviesia.
Austral. J. Bot. 30: 1–9.
De Vattimo I. 1955. Notice sur la tribu Apodantheae R.Br.
(Raffl esiaceae). Taxon 4: 211–212.
De Vattimo I. 1956. Notes on Apodanthes caseariae Poit. and
Pilostyles calliandrae (Gardn.) R.Br. (Raffl esiaceae –
Apodantheae). Not. Syst. 15: 225–229.
De Vattimo I. 1971. Contribução ao conhecimento da tribu
Apodantheae R.Br. Parte 1 – Conspecto das especies
(Raffl esiaceae). Rodriguesia 26: 37–62.
De Vattimo I. 1978. Uma nova especie de Apodanthes Poit.
(Raffl esiaceae). Rodriguesia 29: 269–306.
Emmons LH, J Nias, and A Brium. 1991. The fruit and con-
sumers of Raffl esia keithii (Raffl esiaceae). Biotropica 23:
197–199.
Ernst A and E Schmid. 1913. Über Blüte und Frucht von
Raffl esia. Ann. Jard. Bot. Buitenz. 2, 12: 1–58.
Forstmeier L, F Weberling, and HC Weber. 1983. Zum
Parasitismus von Cytinus hypocistis L. (Raffl esiaceae). Beitr.
Biol. Pfl . 58: 299–311.
68 Subclass I. MAGNOLIIDAE
Garcia Franco JG, V Rico Gray. 1997. Reproductive biology of
the holoparasitic endophyte Bdallophyton bambusarum
(Raffl esiaceae). Bot. J. Linn. Soc. 123(3): 237–247.
Griffi th W. 1845. On the root-parasites referred by authors to
Rhizantheae: and on various plants related to them. Trans.
Linn. Soc. London 19: 303–347.
Gusowska I. 1964. Reinvestigation of embryo sac development,
fertilization, and early embryogeny in Cytinus hypocistis.
Acta Soc. Bot. Polon. 33: 157–166.
Gusowska I. 1966. Microsporogenesis and chromosome number
in Cytinus hypocistis L. Acta Soc. Bot. Polon. 35: 445–454.
Harms H. 1945. Raffl esiaceae. In: A Engler and K Prantl, eds.
Die natürlichen Pfl anzenfamilien, S. 243–281. 2 Aufl .
Leipzig, Berlin.
Hegnauer R. 1997. Phytochemistry of Raffl esiaceae. In:
C Kalkman et al., eds. Flora Malesiana, ser. I, 13: 10,
Leiden.
Heinricher E. 1905. Beiträge zur Kenntnis der Raffl esiaceae: I.
Denkschr. Akad. Wiss. Wien, Math.-Naturw. Kl., 78: 1–25,
57–81.
Heinricher E. 1917. Zur Kenntnis der Blüte von Cytinus hypo-
cistis. Ber. Deutsch. Bot. Ges. 35(3): 513–517.
Heinricher E. 1934. Zur Frage der Artbildung bei Cytinus hypo-
cistis nebst anderen Bemerkungen. Ber. Deutsch. Bot. Ges.
52(I): 48–53.
Hosseus CC. 1907. Eine neue Raffl esiaceengattung aus Siam.
Bot. Jahrb. Syst. 41:55–61.
Justesen PT. 1922. Morphological and biological notes on
Raffl esia fl owers observed in the Highlands of Mid-Sumatra
(Padangsche Bovenlanden). Ann. Jard. Bot. Buitenz. 32:
64–87.
Kuijt J, D Bray, and AR Olson. 1985. Anatomy and ultras truc-
ture of the endophytic system of Pilostyles thurberi
(Raffl esiaceae). Canad. J. Bot. 63: 1231–1240.
Mat Salleh K, A Latiff. 1995. On the morphology of the female
fl ower of Raffl esia tengku-adlinii and notes on the status of
R.borneensis (Raffl esiaceae). Flora Males. Bull. 11(6):
425–428.
Meijer W. 1958. A contribution to the taxonomy and biology of
Raffl esia arnoldi in West Sumatra. Ann. Bogor. 3: 33–44.
Meijer W. 1985. Saving the world’s largest fl ower. Natl. Geogr.
Mag. 168(I): 136–140.
Meijer W. 1993. Raffl esiaceae. In: K Kubitzki, ed. The families
and genera of fl owering plants, vol. 2, pp. 557–563. Springer,
Berlin/Heidelberg/New York.
Meijer W. 1997. Raffl esiaceae. In: C Kalkman et al., eds. Flora
Malesiana, ser. 1, 13: 1–42. Leiden.
Meijer W and JF Veldkamp. 1988. A revision of Rhizanthes
(Raffl esiaceae). Blumea 31: 329–342.
Nickrent DL, A Blarer, Q Yin-Long, R Vidal-Russell, and FE
Anderson. 2004. Phylogenetic inference in Raffl esiales: the
infl uence of rate heterogeneity and horizontal gene transfer.
BMC Evol. Biol. 4: 40 – http://www.biomedcentral.
com/14711–148/4/40
Olah L. 1960. Cytological and morphological investigations in
Raffl esia arnoldi R. Br. Bull. Torrey Bot. Club 87: 406–416.
Ponzi R and P Pizzolongo. 1976. Cytinus hypocistis L.
Embryogenesis: Ultrastructural aspect of megasporo-genesis
and megagametogenesis. J. Submicroscop. Cytol. 8:
327–336.
Ponzi R and P Pizzolongo. 1982. Cytinus hypocistis em-bryo-
genesis: Some biological and ultrastructural aspects of fertil-
ization and embryo development. Nuovo Giorn. Bot. Ital.
116: 149–166.
Royen P van. 1963. Sertulum Papuanum: 8. Raffl esiaceae. Nova
Guinea Bot. 14: 243–245.
Rutherford RJ. 1970. The anatomy and cytology of Pilostyles
thurberi Gray (Raffl esiaceae). Aliso 7: 263–288.
Schaar F. 1898. Über den Bau des Thallus von Raffl esia rochus-
senii Teijsm et Binn. Sitzungsber. K. Akad. Wiss. Wien,
Math.-Naturw. Kl., 107: 1039–1056.
Solms-Laubach HG. 1874a. Über den Thallus von Pilostyles
haussknechtii. Bot. Z. 32: 44–59, 65–74.
Solms-Laubach HG. 1874b. Über den Bau des Samens in den
Familien der Raffl esiaceae und Hydnoraceae. Bot. Z. 32:
337–342, 353–358, 369–374, 385–389.
Solms-Laubach HG. 1875. Das Haustorium der Loranthaceen
und der Thallus der Raffl esiaceen und Balanophoreen.
Abhandl. Naturf. Ges. Halle 13(I): 1–40.
Solms-Laubach HG. 1876. Die Entwicklung der Blüten bei
Brugmansia zippelii und Aristolochia clematitis. Bot. Z. 34:
448–461, 464–470, 480–489,479–503.
Solms-Laubach HG. 1898. Die Entwicklung des Ovulum und
des Samens bei Raffl esia und Brugmansia. Ann. Jard. Bot.
Buitenz., Suppl. 2: 11–22.
Solms-Laubach HG. 1901. Raffl esiaceae. In: A Engler und K
Prantl, eds. Das Pfl anzenreich. IV, 75: 1–19. Berlin.
Takhtajan AL, NR Meyer, and VN Kosenko. 1985. Pollen mor-
phology and classifi cation in Raffl esiaceae s. 1. Bot. Zhurn.
70: 153–162 (in Russian).
Terekhin ES. 1988. Raffl esiales. In: A Takhtajan, ed. Comparative
seed anatomy, vol. 2, pp. 114–119. Nauka, Leningrad (in
Russian).
Terekhin ES and GM Anisimova. 1981. Raffl esiaceae. In: MS
Yakovlev, ed. Comparative embryology of fl owering plants,
pp. 96–100. Nauka, Leningrad (in Russian).
Winlder H. 1927. Über eine Raffl esia aus Zentralborneo. Planta
4: 1–79.
A. Takhtajan, Flowering Plants, 69
© Springer Science + Business Media B.V. 2009
Mostly herbaceous plants. Parenchymatous tissues
usually without ethereal cells. Stomata mostly anomo-
cytic. Vessels usually with simple perforations. Sieve-
element plastids of S-type. Flowers actinomorphic or
zygomorphic, bisexual or less often unisexual, spiro-
cyclic or more often cyclic. Tapetum secretory, rarely
amoeboid (Kingdonia and some species of Mahonia
and Berberis). Microsporogenesis simultaneous or
rarely successive (some species of Mahonia). Pollen
grains 2-celled or 3-celled, 3-colpate or of derived
types. Gynoecium apocarpous, syncarpous, or paracar-
pous. Ovules mostly anatropous, bitegmic or sometimes
unitegmic, crassinucellate or seldom tenuinucellate.
Female gametophyte commonly of Polygonum-type.
Endosperm cellular or more often nuclear. Fruits of
various types. Seeds with small embryo and copious
endosperm or less often with large embryo and little or
no endosperm. Commonly producing diverse kinds of
isoquinoline alkaloids, but without ellagic acid and
iridoid compounds.
Most probably derived from some archaic
Magnoliidae.
Bibliography
Barthlott W and I Theisen. 1995. Epicuticular wax ultrastructure
and classifi cation of Ranunculifl orae. In: U Jensen and
JW Kadereit, eds. Systematics and evolution of the
Ranunculifl orae. Plant Syst. Evol., Suppl. 9: 39–45.
Behnke H-D 1971. Sieve-tube plastids in Magnoliidae and
Ranunculidae in relation to systematics. Taxon 20:
723–730.
Behnke H-D. 1995. Sieve-element plastids, phloem proteins,
and the evolution of the Ranunculanae. In: U Jensen and
JW Kadereit, eds. Systematics and evolution of the
Ranunculifl orae. Plant Syst Evol., Suppl. 9: 25–37.
Blackmore S, P Stafford, and V Persson. 1995. Palynology and
systematics of Ranunculifl orae. In: U Jensen and JW Kadereit,
eds. Systematics and evolution of the Ranunculifl orae. Plant
Syst. Evol., Suppl. 9: 71–82.
Brückner C. 2000. Clarifi cation of the carpel. Number in
Papaverales, Capparales, and Berberidaceae. Bot. Rev. 66:
155–309.
Carlquist S. 1995. Wood anatomy of Ranunculifl orae: a sum-
mary. In: U Jensen and JW Kadereit, eds. Systematics and
evolution of the Ranunculifl orae. Plant Syst. Evol., Suppl. 9:
11–24.
Endress PK. 1995. Floral structure and evolution in Ranunculanae.
In: U Jensen and JW Kadereit, eds. Systematics and evolu-
tion of the Ranunculifl orae. Plant Syst. Evol., Suppl. 9:
47–61.
Hiepko P. 1965. Vergleichend-morphologische und entwick-
lungsgeschichtliche Untersuchungen über das Perianth bei
den Polycarpicae. Bot. Jahrb. Syst. 84: 359–508.
Hoot SB and PR Crane. 1995. Inter-familial relationships in the
Ranunculidae based on molecular systematics. In: U Jensen
and JW Kadereit, eds. Systematics and evolution of the
Ranunculifl orae. Plant Syst. Evol., Suppl. 9: 119–131.
Jensen U. 1995. Secondary compounds of the Ranunculifl orae.
Plant Syst. Evol., Suppl. 9: 85–97.
Jensen U and JW Kadereit, eds. 1995. Systematics and evolution
of the Ranunculifl orae. Plant Syst. Evol., Suppl. 9:
133–145.
Kubitzki K. 1995. Ranunculifl orae – delimitation, phylogeny,
diversifi cation. In: U Jensen and JW Kadereit, eds.
Systematics and evolution of the Ranunculifl orae. Plant Syst.
Evol., Suppl. 9: 1–10.
Loconte H, LM Campbell, and SW Stevenson. 1995. Original
and familial relationships of the Ranunculifl orae. In:
U Jensen and JW Kadereit, eds. Systematics and evolution of
the Ranunculifl orae. Plant Syst. Evol., Suppl. 9: 99–118.
Smith AC. 1971 (1972). An appraisal of the orders and families
of primitive extant angiosperms. J. Indian Bot. Soc. (Golden
Jubilee) 50A: 215–226.
Walker JW. 1976. Comparative pollen morphology and phylog-
eny of the ranalean complex, pp. 241–299. In: CB Beck, ed.
Origin and early evolution of angiosperms. New York.
Subclass II. RANUNCULIDAE
70 Subclass II. RANUNCULIDAE
Superorder PROTEANAE
Order 18. PLATANALES
Large, monoecious, deciduous trees, often with mottled
bark falling off in large fl akes; hairs candelabriform.
Vessels with scalariform and simple perforations; ves-
sels without vestured pits. Axial parenchyma apotra-
cheal, diffuse or in uniseriate bands. Rays homogeneous.
Sieve-element plastids of S-type. Nodes multilacu-
nar (7). Leaves alternate, medium-sized to large, pal-
mately veined and lobed or seldom (Platanus kerrii)
pinnately veined and entire. Leaves without a persistent
basal meristem (Watson and Dallwitz 2006). The peti-
ole usually mitriform at the base and enclosing the axil-
lary bud or seldom (P. kerrii) buds free, teeth glandular,
with a cavity. Candlabriform trichomes with whorls of
arms cover the abaxial surface of leaves (Carpenter
et al. 2005). Stipules concrescent (around the stem),
ochreate, scaly, caducous. Stomata anomocytic, latero-
cytic or paracytic. Infl orescences monoecious, long
pedunculate, pendulous, of 1 or more globular heads of
numerous fl owers, each head subtended by a circular
bract, smaller bracts interspersed among the fl owers.
Individual fl owers small, inconspicuous, basically
4-merous; sepals 3-4(-7) free or basally connate, not
vascularized; corolla in male fl owers 3-4(-7), tiny or
vestigial. Stamens as many as and opposite the sepals;
fi laments very short. Anthers basifi xed or adnate,
latrorse, 2-locular, tetrasporangiate, with a connective
enlarged at the apex into a peltate appendage.
Microsporogenesis simultaneous. Tapetum secretory.
Pollen grains 2-celled, 3-colpate or 6-rugate. Male fl ow-
ers sometimes with vestigial carpels. Female fl owers
with 3–4 staminodia, or without staminodes. Gynoecium
apocarpous, of 5–8 or less often 3–4 or 9 carpels in 2–3
cycles; stylodia long, linear, with decurrent stigma; car-
pels are not completely sealed distally. Ovules orthotro-
pous or slightly hemitropous, usually solitary, rarely
two, pendulous, bitegmic, crassinucellate. Placentation
apical to marginal. Endosperm nuclear. Fruits of
achenes, rarely of almost follicles, with tuft of basal
hairs. Seeds small, with thin testa and tegmen com-
pletely disappearing in the ripe seed; embryo straight,
rather large; endosperm scanty. Contain cyanogenic
(cynogenic constituents tyrosine-derived), proanthocy-
anidins (cyanidin and delphinidin), fl avonols (kaemp-
ferol, quercetin, and myricetin), n = 7, 8, 16–21.
1. PLATANACEAE
T.G. Lestiboudois 1826. 1/10. From the Balkan
Peninsula to western Himalayas, Indochina, and
America from Canada to Mexico.
Platanus.
Bibliography
Baas P. 1969. Comparative anatomy of Platanus kerrii Gagenep.
Bot. J. Linn. Soc. 62: 413–421.
Boothroyd LE. 1930. The morphology and anatomy of the infl o-
rescence and fl ower of the Platanaceae. Am. J. Bot. 17:
678–693.
Brett DW. 1979. Ontogeny and classifi cation of the stomatal
complex of Platanus L. Ann. Bot. 44: 249–251.
Bretzler E. 1938. Bau der Platanenblüte und systematische
Stellung der Platanen. Bot. Jahrb. Syst. 62: 305–309.
Carpenter RJ, RS Hill, and GJ Jordan. 2005. Leaf cuticular mor-
phology links Platanaceae and Proteaceae. Int. J. Plant Sci.
166: 843–855.
Ernst WR. 1963. The genera of Hamamelidaceae and Platanaceae in
the southeastern United States. J. Arnold Arbor. 44: 193–210.
Feng Y, S-H Oh, and PS Manos. 2005. Phylogeny and historical
biogeography of the genus Platanus as inferred from nuclear
and chloroplast DNA. Syst. Bot. 30: 786–799.
Floyd SK, VT Lerner, and WE Friedman. 1999. A developmen-
tal and evolutionary analysis of embryology in Platanus
(Platanaceae), a basal eudicot. Am. J. Bot. 86: 1523–1537.
Hsiao J-Y. 1973. A numerical taxonomic study of the genus
Platanus based on morphological and phenolic characters.
Am. J. Bot. 60: 678–684.
Kubitzki K. 1993. Platanaceae. In: K Kubitzki, ed. The families
and genera of vascular plants, 2: 521–522. Springer, Berlin/
Heidelberg/New York.
Leroy J-F. 1982. Origine et evolution du genre Platanus
(Platanaceae). C. R. Acad. Sci. Paris, ser. 3, 295: 251–254.
Maslova NP. 2003. Extinct and extant Platanaceae and
Hamamelidaceae: morphology, systematics, and phylogeny.
Paleontol. J. 37: 467–590.
Mindell RA, RA Stockey, and G Beard. 2006. Anatomically pre-
served staminate infl orescences of Gynoplatananthus oyster-
bayensis gen. et sp. nov. (Platanaceae) and associated
pistillate frutifi cations from the Eocene of Vancouver Island,
British Columbia. Int. J. Plant Sci. 167: 591–600.
Oginuma K and H Tobe. 1991. Karyomorphology and evolution
in some Hamamelidaceae and Platanaceae (Hamamelididae:
Hamamelidales). Bot. Mag. Tokyo 104: 115–135.
Rieseberg LH and DE Soltis. 1987. Flavonoids of fossil Miocene
Platanus and its extant relatives. Biochem. Syst. Ecol. 15:
109–112.
Schwarzwalder RN. 1986. Systematics and early evolution of
the Platanaceae. Ph.D. dissert. Indiana University.
Schwarzwalder RN and DL Dilcher. 1991. Systematic place-
ment of the Platanaceae in the Hamamelididae. Ann.
Missouri Bot. Gard. 78: 962–969.