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ARISTOLOCHIACEAE
Review of Palaeobotany and Palynology 123 (2003) 47^55 www.elsevier.com/locate/revpalbo
The Northwest European Pollen Flora, 62
ARISTOLOCHIACEAE Ch. Mulder Laboratory of Palaeobotany and Palynology, Utrecht University, Utrecht, The Netherlands
Literature Agababjan (1973), Andrew (1984), Bertsch (1942), Beug (1961), Erdtman (1952), Heusser (1971), Kuprianova and Alyoshina (1972), Mitroiu (1970), Reille (1992, 1995, 1998), Roure (1985), Stachurska et al. (1981), Straka (1981), Tarnavschi et al. (1981), Valde¤s et al. (1987), Walker (1974, 1976), Wodehouse (1935).
Introduction The Aristolochiaceae is a small family with only seven genera and some 625 species (Heywood, 1993). Their distribution is mainly pantropical, with the 20 temperate European taxa growing chie£y in the Mediterranean region. The genus Aristolochia is the largest with about 300 species, only three of which are native in Northwest Europe (Jalas and Suominen, 1976; Tutin et al., 1993). The second genus of importance, Asarum, is represented by A. europaeum. The pollen grains of the Aristolochiaceae have been neglected in most palynological works, or they were considered in some keys only as rare grains (like A. europaeum in Moore et al., 1991). This appears to be due to the low pollen production and dispersal of Aristolochiaceae. The family is eurypalynous, with a great diversity of sexine sculpturing types and pollen classes (Nair, 1970). For instance, Aristolochia pollen grains have been described as verrucate (Heusser, 1971; Roubik and Moreno, 1991), verrucate-areolate (El Ghazali, * Present address: National Institute for Public Health and the Environment, P.O. Box 1, 3720 BA Bilthoven, The Netherlands. E-mail address: [email protected] (Ch. Mulder).
1993), verrucate-reticulate (Mitroiu, 1970), scabrate-reticulate (Huang, 1972), ¢nely reticulate (Johri and Bhatnagar, 1955; Rao and Ong, 1974), or psilate (Bertsch, 1942; Beug, 1961). Literature of pertinent palynological records is scarce although pollen of Aristolochiaceae has been recently recorded in present-day moss samples (surface samples) along the Dutch coast (Aristolochia in Mulder, 1996). The terminology used follows that recommended by Punt et al. (1994).
Specimens examined All the specimens listed here are housed in the Herbarium of Utrecht (U) unless otherwise stated. Aristolochia clematitis L. ^ England: Jackson and Druce 1455 (BM); France: Campbell s.n. (BM), Exc. Biol. Stud. Utrecht 32-295; Italy: Ka«ssner 55008 (J); The Netherlands: Hekking s.n., Fresh Material Punt Anno 1961. A. longa L. = A. paucinervis Pomel p.p. A. paucinervis Pomel ^ Italy: Ka«ssner 55012 (J); Portugal: Kostermans and Kruyt 36-49; van der Knaap and van Leeuwen 89-4; Spain: Alston 10373 (BM). A. rotunda L. ^ France: Hekking 58-12A, Brijl 63-35; Italy: Cook 83-4046, Davis and Sutton 64005 (BM), Ka«ssner 55013 (J), Fresh Material Mulder Anno 1994. Asarum europaeum L. ^ Austria: Kramer and Westra 67-4139;
0034-6667 / 02 / $ ^ see front matter I 2002 Elsevier Science B.V. All rights reserved. PII: S 0 0 3 4 - 6 6 6 7 ( 0 2 ) 0 0 1 5 3 - 7
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C. Mulder / Review of Palaeobotany and Palynology 123 (2003) 47^55
Czechoslovakia: Tr›ebova¤ 59-8220; France: Desplantes 19195 (BM), Exc. Biol. Stud. Utrecht 49-449; Germany: Fresh Material Mulder 1995, Rutters s.n.; Switzerland: Kramer 47-70, Mennega s.n.
Key to the pollen types 1.a. Tectum with gemmate processes; columellae indistinct ....................... Asarum europaeum type b. Tectum without gemmate processes; distinct columellae................ Aristolochia clematitis type
Description of pollen types Aristolochia clematitis type (Plates 1^3) Pollen class: Inaperturate. Shape: Spheroidal to irregularly ellipsoidal. Apertures : Absent. Exine: Fairly thin to thick. Nexine about as thick as sexine. Sexine 1 of distinct columellae, uniform in thickness. Sexine 2 a perforate tectum ; tectum extremely thin often undulating. Sexine 3 absent. Ornamentation: Tectate, microperforate. Puncta more or less circular, irregularly distributed. Surface slightly rugulate. Columellae distinct, irregular in outline, irregularly arranged, crowded. Outlines : More or less circular or weakly angular, often £attened. Measurements: Glycerine jelly ^ Largest diameter 35.0^61.0 Wm. Exine 1.5^4.0 Wm. Tectum less than 1 Wm. Silicone oil ^ Largest diameter 32.0^48.0 Wm. Species: Aristolochia clematitis, A. paucinervis, A. rotunda Key to the species and groups (use U1000 magni¢cation) 1.a. Exine thinner than 2.5 Wm, ornamentation slightly scabrate............... Aristolochia clematitis
Aristolochia clematitis A. paucinervis A. rotunda
b. Exine thicker than 2.5 Wm, ornamentation verrucate .................. Aristolochia rotunda group (Aristolochia paucinervis, A. rotunda) Comments The Aristolochia clematitis type is characterised by its absence of apertures and undulating tectum. Pollination of the investigated species of Aristolochia is similar in all species (Solereder, 1889). The tendency to form coated clumps of pollen grains connected by non-sporopollenin viscous ¢bres appears throughout the genus (Hesse, 1986). Pollen of Aristolochia clematitis has a thin sexine, whereas the exine of the Aristolochia rotunda group is signi¢cantly thicker. The exine of all species often splits into two halves. The splitting of the thin area in A. clematitis pollen grains was already noted by Andrew (1984). All pollen grains larger than 40 Wm in diameter in the specimens investigated (very few of the grains above 50 Wm) belong to the Aristolochia rotunda group; because of the overlapping sizes the smaller grains can belong to both groups. The description of the A. rotunda group corresponds to the Aristolochia paucinervis type of Valde¤s et al. (1987). Asarum europaeum type (Plate 4) Pollen class: Faintly 3-porate, often indistinctly. Shape: Spheroidal to slightly ellipsoidal. Apertures: Porus-like, often indistinct; margins irregular. Exine: Fairly thin. Sexine about as thick as nexine. Sexine 1 of short columellae. Sexine 2 a thin tectum. Sexine 3 of distinct verrucae or gemmae. Ornamentation : Tectate, microperforate, verrucate or gemmate. Columellae distinct, more or less circular in outline, regularly arranged, more or less spaced. Outlines: Circular to slightly angular. Measurements: Glycerine jelly ^ Largest diameter
Glycerine jelly (Wm)
Silicone oil (Wm)
Exine (Wm)
26.0^(34.5)^40.0 33.0^(42.0)^55.0 28.0^(40.5)^48.0
25.0^(35.0)^40.0 26.0^(39.5)^52.0 20.0^(35.5)^48.0
1.7 3.3 2.9
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26.0^(36.5)^45.0 Wm. Silicone oil ^ Largest diameter 40.0^(46.0)^51.0 Wm. Exine about 1 Wm. Diameter verrucae and gemmae 1.5 Wm or less. Species: Asarum europaeum Comments The Asarum europaeum type is characterised by its irregular porus-like apertures and the verrucate-gemmate ornamentation. These characters clearly di¡ers this type from the Aristolochia clematitis type. In the literature pollen of Asarum is usually described as inaperturate with a gemmate or baculate-verrucate ornamentation (Straka, 1981). Some authors describe Asarum pollen as inaperturate or 3-zonocolpate (Ikuse, 1956; Mi and Yang, 1991). Reille (1992) includes A. europaeum in both pollen classes, viz. inaperturate and 3-zonocolpate.
References Agababjan, V.Sh., 1973. Pollen of primitive Angiospermae (in Russian). Bot. Inst. Akad. Nauk Armen. SSR, 169 pp., 28 pl. Andrew, R., 1984. A Practical Pollen Guide to the British Flora. Technical Guide 1. Quaternary Research Association, Cambridge, 139 pp. Bertsch, K., 1942. Lehrbuch der Pollenanalyse. F. Enke, Stuttgart, 194 pp. Beug, H.-J., 1961. Leitfaden der Pollenbestimmung fu«r Mitteleuropa und angrenzende Gebiete. Gustav Fischer Verlag, Jena, 63 pp. El Ghazali, G.E.B., 1993. A study on the pollen £ora of Sudan. Rev. Palaeobot. Palynol. 76, 95^345. Erdtman, G., 1952. Pollen Morphology and Plant Taxonomy. Angiosperms. Almqvist and Wiksell, Stockholm, 539 pp. Hesse, M., 1986. Nature, form and function of pollen-connecting threads in angiosperms. In: Blackmore, S., Ferguson, I.K. (Eds.), Pollen and Spores: Form and Function. Academic Press, London, pp. 109^118. Heusser, C.J., 1971. Pollen and Spores of Chile. Modern Types of the Pteridophyta, Gymnospermae, and Angiospermae. The University of Arizona Press, 167 pp. Heywood, V.H., 1993. Flowering Plants of the World. B.T. Batsford, London, 336 pp. Huang, T.C., 1972. Pollen Flora of Taiwan. National Taiwan University, Botany Department Press, 297 pp. Ikuse, M., 1956. Pollen Grains of Japan (in Japanese). Hirokawa, Tokio, 303 pp. Jalas, J., Suominen, J., 1976. Aristolochiaceae. In: Atlas Florae Europaeae, 3: Salicaceae to Balanophoraceae. The Com-
49
mittee for Mapping the Flora of Europe and Societas Biologica Fennica Vanamo, pp. 115^120. Johri, B.M., Bhatnagar, S.P., 1955. A contribution to the morphology and life-history of Aristolochia. Phytomorphology 5, 122^137. Kuprianova, L.A., Alyoshina, L.A., 1972. Pollen and Spores of Plants from the Flora of European Part of the USSR, I (in Russian). The Academy of Science of the USSR/The Komarov Botanical Institute, 171 pp. Mi, Q.W., Yang, C.S., 1991. Pollen morphology of Asarum in China (in Chinese). Acta Phytotaxon. Sin. 29, 164^171. Ł tudes morphopolliniques et des aspects Mitroiu, N., 1970. E embryologiques sur les ‘Polycarpicae’ et Helobiae, avec des conside¤rations phyloge¤ne¤tiques. Acta Bot. Horti Bucur., 243 pp. Moore, P.D., Webb, J.A., Collinson, M.E., 1991. Pollen Analysis, 2nd edn. Blackwell Scienti¢c, Oxford, 216 pp. Mulder, Ch., 1996. Analisi comparata della struttura della vegetazione attuale con lo studio palynologico di campioni di super¢cie in di¡erenti regioni europee. PhD Bot. Sci. (VIII Cycle) Nat. Libr. Rome Florence, 160 pp. Nair, P.K.K., 1970. Pollen Morphology of Angiosperms. A Historical and Phylogenetic Study. Lucknow Publishing House, 160 pp. Punt, W., Blackmore, S., Nilsson, S., Le Thomas, A., 1994. Glossary of pollen and spore terminology. LPP Contr. Ser., 1, Utrecht, 71 pp. Rao, A.N., Ong, E.T., 1974. Pollen morphology of certain tropical plants. J. Palynol. 10, 1^37. Reille, M., 1992. Pollen et Spores d’Europe et d’Afrique du Nord. Lab. Bot. Hist. Palynol., Marseille, 520 pp. Reille, M., 1995. Pollen et Spores d’Europe et d’Afrique du Nord. Suppl. 1. Lab. Bot. Hist. Palynol., Marseille, 327 pp. Reille, M., 1998. Pollen et Spores d’Europe et d’Afrique du Nord. Suppl. 2. Lab. Bot. Hist. Palynol., Marseille, 521 pp. Roubik, D.W., Moreno, J.E., 1991. Pollen and Spores of Barro Colorado Island. Monographs in Systematic Botany from the Missouri Botanical Garden 36, 1^270. Roure, J.M., 1985. Palinolog|¤a Ibe¤rica. Fam. 1 a 20. Cupressaceae a Betulaceae. Orsis 1, 43^69. Solereder, H., 1889. Aristolochiaceae. In: Engler, A., Prantl, K. (Eds.). Die natu«rlichen P£anzenfamilien, III Teil, 1. Abt. Verlag Wilhelm Engelmann, pp. 264^273. Stachurska, A., Sadowska, A., Kuszell, T., 1981. Kartoteka Palinologiczna Roslin Polskich. Opol. Tow. Przyj. Nauk Zesz. Przyr. 21, 257^266. Straka, H., 1981. Zur Pollenkunde (Palynologie) der in Band III/1 behandelten Familien. In: Hegi, G. (Ed.), Illustrierte Flora von Mittel-Europa. Band III. Teil 1, 3rd edn. (Conert, H.J., Hamann, U., Schultze-Motel, W., Wagenitz, G., Eds.). Verlag Paul Parey, Berlin, pp. 488^504. Tarnavschi, I.T., SVerba›nescu-Jitariu, G., Mitroiu-Ra›dulescu, N. and Ra›dulescu, D., 1981. A Monograph of Pollen in the Flora of Rumania. I (in Rumanian). Editura Academiei Roma›ne, Bucarest, 144 pp. Tutin, T.G., Burges, N.A., Chater, A.O., Edmondson, J.R., Heywood, V.H., Moore, D.M., Valentine, D.H., Walters,
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S.M., Webb, D.A. (Eds.), 1993. Flora Europaea, Volume 1: Psilotaceae to Platanaceae (2nd ed.). Cambridge University, pp. 87^89. Valde¤s, B., D|¤ez, M.J., Ferna¤ndez, I. 1987. Atlas polinico de Andalucia Occidental. Inst. Desarro. Reg. No. 43, Univ. Sevilla Excma. Dip. Cadiz, 450 pp. Walker, J.W., 1974. Aperture evolution in the pollen of primitive Angiosperms. Am. J. Bot. 61 (10), 1112^1137.
Walker, J.W., 1976. Evolutionary signi¢cance of the exine in the pollen of primitive angiosperms. In: Ferguson, I.K., Muller, J. (Eds.), The Evolutionary Signi¢cance of the Exine. Linnean Press, pp. 251^308. Wodehouse, R.P., 1935. Pollen Grains. McGraw-Hill, New York, 574 pp.
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Plate descriptions (all ¢gures U2000 except where otherwise stated) PLATE 1 (p. 52) Aristolochia clematitis L. (Campbell s.n.) 1. SEM micrograph; overall view. 2. SEM micrograph; ornamentation (U5000). Aristolochia paucinervis Pomel (Alston 10373) 3. SEM micrograph; overall view. 4. SEM micrograph; ornamentation (U5000). PLATE 2 (p. 53) Aristolochia paucinervis Pomel (¢g. 1, Alston 10373, ¢gs. 2^4, Van der Knaap and Van Leeuwen 89-4) 1. SEM micrograph; cross-section of the exine (U5000). 2. Cross-section. 3. Ornamentation at high focus. 4. Ornamentation at low focus. Aristolochia rotunda L. (Hekking 58-12A) 5. Cross-section. PLATE 3 (p. 54) Aristolochia rotunda L. (¢gs. 1^2, Davis and Sutton 64005; ¢gs. 3^4, Hekking 58-12A) 1. SEM micrograph; overall view. 2. SEM micrograph; ornamentation (U5000). 3. Ornamentation at high focus. 4. Ornamentation at low focus. PLATE 4 (p. 55) Asarum europaeum L. (¢gs. 1, Desplantes 19195; ¢gs. 2^5, Fresh Material Mulder Anno 1995) 1. SEM micrograph; polar view. 2. Polar view; ornamentation at high focus. 3. Polar view; ornamentation at medium focus. 4. Polar view; ornamentation at low focus. 5. Polar view, cross-section.
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PLATE 1 (Aristolochia clematitis type: Aristolochia clematitis, 1^2; Aristolochia paucinervis, 3^4)
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PLATE 2 (Aristolochia clematitis type: Aristolochia paucinervis, 1^4; Aristolochia rotunda, 5)
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PLATE 3 (Aristolochia clematitis type: Aristolochia rotunda)
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PLATE 4 (Asarum europaeum type: Asarum europaeum)
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The Northwest European Pollen Flora, 62
ARISTOLOCHIACEAE Ch. Mulder Laboratory of Palaeobotany and Palynology, Utrecht University, Utrecht, The Netherlands
Literature Agababjan (1973), Andrew (1984), Bertsch (1942), Beug (1961), Erdtman (1952), Heusser (1971), Kuprianova and Alyoshina (1972), Mitroiu (1970), Reille (1992, 1995, 1998), Roure (1985), Stachurska et al. (1981), Straka (1981), Tarnavschi et al. (1981), Valde¤s et al. (1987), Walker (1974, 1976), Wodehouse (1935).
Introduction The Aristolochiaceae is a small family with only seven genera and some 625 species (Heywood, 1993). Their distribution is mainly pantropical, with the 20 temperate European taxa growing chie£y in the Mediterranean region. The genus Aristolochia is the largest with about 300 species, only three of which are native in Northwest Europe (Jalas and Suominen, 1976; Tutin et al., 1993). The second genus of importance, Asarum, is represented by A. europaeum. The pollen grains of the Aristolochiaceae have been neglected in most palynological works, or they were considered in some keys only as rare grains (like A. europaeum in Moore et al., 1991). This appears to be due to the low pollen production and dispersal of Aristolochiaceae. The family is eurypalynous, with a great diversity of sexine sculpturing types and pollen classes (Nair, 1970). For instance, Aristolochia pollen grains have been described as verrucate (Heusser, 1971; Roubik and Moreno, 1991), verrucate-areolate (El Ghazali, * Present address: National Institute for Public Health and the Environment, P.O. Box 1, 3720 BA Bilthoven, The Netherlands. E-mail address: [email protected] (Ch. Mulder).
1993), verrucate-reticulate (Mitroiu, 1970), scabrate-reticulate (Huang, 1972), ¢nely reticulate (Johri and Bhatnagar, 1955; Rao and Ong, 1974), or psilate (Bertsch, 1942; Beug, 1961). Literature of pertinent palynological records is scarce although pollen of Aristolochiaceae has been recently recorded in present-day moss samples (surface samples) along the Dutch coast (Aristolochia in Mulder, 1996). The terminology used follows that recommended by Punt et al. (1994).
Specimens examined All the specimens listed here are housed in the Herbarium of Utrecht (U) unless otherwise stated. Aristolochia clematitis L. ^ England: Jackson and Druce 1455 (BM); France: Campbell s.n. (BM), Exc. Biol. Stud. Utrecht 32-295; Italy: Ka«ssner 55008 (J); The Netherlands: Hekking s.n., Fresh Material Punt Anno 1961. A. longa L. = A. paucinervis Pomel p.p. A. paucinervis Pomel ^ Italy: Ka«ssner 55012 (J); Portugal: Kostermans and Kruyt 36-49; van der Knaap and van Leeuwen 89-4; Spain: Alston 10373 (BM). A. rotunda L. ^ France: Hekking 58-12A, Brijl 63-35; Italy: Cook 83-4046, Davis and Sutton 64005 (BM), Ka«ssner 55013 (J), Fresh Material Mulder Anno 1994. Asarum europaeum L. ^ Austria: Kramer and Westra 67-4139;
0034-6667 / 02 / $ ^ see front matter I 2002 Elsevier Science B.V. All rights reserved. PII: S 0 0 3 4 - 6 6 6 7 ( 0 2 ) 0 0 1 5 3 - 7
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C. Mulder / Review of Palaeobotany and Palynology 123 (2003) 47^55
Czechoslovakia: Tr›ebova¤ 59-8220; France: Desplantes 19195 (BM), Exc. Biol. Stud. Utrecht 49-449; Germany: Fresh Material Mulder 1995, Rutters s.n.; Switzerland: Kramer 47-70, Mennega s.n.
Key to the pollen types 1.a. Tectum with gemmate processes; columellae indistinct ....................... Asarum europaeum type b. Tectum without gemmate processes; distinct columellae................ Aristolochia clematitis type
Description of pollen types Aristolochia clematitis type (Plates 1^3) Pollen class: Inaperturate. Shape: Spheroidal to irregularly ellipsoidal. Apertures : Absent. Exine: Fairly thin to thick. Nexine about as thick as sexine. Sexine 1 of distinct columellae, uniform in thickness. Sexine 2 a perforate tectum ; tectum extremely thin often undulating. Sexine 3 absent. Ornamentation: Tectate, microperforate. Puncta more or less circular, irregularly distributed. Surface slightly rugulate. Columellae distinct, irregular in outline, irregularly arranged, crowded. Outlines : More or less circular or weakly angular, often £attened. Measurements: Glycerine jelly ^ Largest diameter 35.0^61.0 Wm. Exine 1.5^4.0 Wm. Tectum less than 1 Wm. Silicone oil ^ Largest diameter 32.0^48.0 Wm. Species: Aristolochia clematitis, A. paucinervis, A. rotunda Key to the species and groups (use U1000 magni¢cation) 1.a. Exine thinner than 2.5 Wm, ornamentation slightly scabrate............... Aristolochia clematitis
Aristolochia clematitis A. paucinervis A. rotunda
b. Exine thicker than 2.5 Wm, ornamentation verrucate .................. Aristolochia rotunda group (Aristolochia paucinervis, A. rotunda) Comments The Aristolochia clematitis type is characterised by its absence of apertures and undulating tectum. Pollination of the investigated species of Aristolochia is similar in all species (Solereder, 1889). The tendency to form coated clumps of pollen grains connected by non-sporopollenin viscous ¢bres appears throughout the genus (Hesse, 1986). Pollen of Aristolochia clematitis has a thin sexine, whereas the exine of the Aristolochia rotunda group is signi¢cantly thicker. The exine of all species often splits into two halves. The splitting of the thin area in A. clematitis pollen grains was already noted by Andrew (1984). All pollen grains larger than 40 Wm in diameter in the specimens investigated (very few of the grains above 50 Wm) belong to the Aristolochia rotunda group; because of the overlapping sizes the smaller grains can belong to both groups. The description of the A. rotunda group corresponds to the Aristolochia paucinervis type of Valde¤s et al. (1987). Asarum europaeum type (Plate 4) Pollen class: Faintly 3-porate, often indistinctly. Shape: Spheroidal to slightly ellipsoidal. Apertures: Porus-like, often indistinct; margins irregular. Exine: Fairly thin. Sexine about as thick as nexine. Sexine 1 of short columellae. Sexine 2 a thin tectum. Sexine 3 of distinct verrucae or gemmae. Ornamentation : Tectate, microperforate, verrucate or gemmate. Columellae distinct, more or less circular in outline, regularly arranged, more or less spaced. Outlines: Circular to slightly angular. Measurements: Glycerine jelly ^ Largest diameter
Glycerine jelly (Wm)
Silicone oil (Wm)
Exine (Wm)
26.0^(34.5)^40.0 33.0^(42.0)^55.0 28.0^(40.5)^48.0
25.0^(35.0)^40.0 26.0^(39.5)^52.0 20.0^(35.5)^48.0
1.7 3.3 2.9
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26.0^(36.5)^45.0 Wm. Silicone oil ^ Largest diameter 40.0^(46.0)^51.0 Wm. Exine about 1 Wm. Diameter verrucae and gemmae 1.5 Wm or less. Species: Asarum europaeum Comments The Asarum europaeum type is characterised by its irregular porus-like apertures and the verrucate-gemmate ornamentation. These characters clearly di¡ers this type from the Aristolochia clematitis type. In the literature pollen of Asarum is usually described as inaperturate with a gemmate or baculate-verrucate ornamentation (Straka, 1981). Some authors describe Asarum pollen as inaperturate or 3-zonocolpate (Ikuse, 1956; Mi and Yang, 1991). Reille (1992) includes A. europaeum in both pollen classes, viz. inaperturate and 3-zonocolpate.
References Agababjan, V.Sh., 1973. Pollen of primitive Angiospermae (in Russian). Bot. Inst. Akad. Nauk Armen. SSR, 169 pp., 28 pl. Andrew, R., 1984. A Practical Pollen Guide to the British Flora. Technical Guide 1. Quaternary Research Association, Cambridge, 139 pp. Bertsch, K., 1942. Lehrbuch der Pollenanalyse. F. Enke, Stuttgart, 194 pp. Beug, H.-J., 1961. Leitfaden der Pollenbestimmung fu«r Mitteleuropa und angrenzende Gebiete. Gustav Fischer Verlag, Jena, 63 pp. El Ghazali, G.E.B., 1993. A study on the pollen £ora of Sudan. Rev. Palaeobot. Palynol. 76, 95^345. Erdtman, G., 1952. Pollen Morphology and Plant Taxonomy. Angiosperms. Almqvist and Wiksell, Stockholm, 539 pp. Hesse, M., 1986. Nature, form and function of pollen-connecting threads in angiosperms. In: Blackmore, S., Ferguson, I.K. (Eds.), Pollen and Spores: Form and Function. Academic Press, London, pp. 109^118. Heusser, C.J., 1971. Pollen and Spores of Chile. Modern Types of the Pteridophyta, Gymnospermae, and Angiospermae. The University of Arizona Press, 167 pp. Heywood, V.H., 1993. Flowering Plants of the World. B.T. Batsford, London, 336 pp. Huang, T.C., 1972. Pollen Flora of Taiwan. National Taiwan University, Botany Department Press, 297 pp. Ikuse, M., 1956. Pollen Grains of Japan (in Japanese). Hirokawa, Tokio, 303 pp. Jalas, J., Suominen, J., 1976. Aristolochiaceae. In: Atlas Florae Europaeae, 3: Salicaceae to Balanophoraceae. The Com-
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mittee for Mapping the Flora of Europe and Societas Biologica Fennica Vanamo, pp. 115^120. Johri, B.M., Bhatnagar, S.P., 1955. A contribution to the morphology and life-history of Aristolochia. Phytomorphology 5, 122^137. Kuprianova, L.A., Alyoshina, L.A., 1972. Pollen and Spores of Plants from the Flora of European Part of the USSR, I (in Russian). The Academy of Science of the USSR/The Komarov Botanical Institute, 171 pp. Mi, Q.W., Yang, C.S., 1991. Pollen morphology of Asarum in China (in Chinese). Acta Phytotaxon. Sin. 29, 164^171. Ł tudes morphopolliniques et des aspects Mitroiu, N., 1970. E embryologiques sur les ‘Polycarpicae’ et Helobiae, avec des conside¤rations phyloge¤ne¤tiques. Acta Bot. Horti Bucur., 243 pp. Moore, P.D., Webb, J.A., Collinson, M.E., 1991. Pollen Analysis, 2nd edn. Blackwell Scienti¢c, Oxford, 216 pp. Mulder, Ch., 1996. Analisi comparata della struttura della vegetazione attuale con lo studio palynologico di campioni di super¢cie in di¡erenti regioni europee. PhD Bot. Sci. (VIII Cycle) Nat. Libr. Rome Florence, 160 pp. Nair, P.K.K., 1970. Pollen Morphology of Angiosperms. A Historical and Phylogenetic Study. Lucknow Publishing House, 160 pp. Punt, W., Blackmore, S., Nilsson, S., Le Thomas, A., 1994. Glossary of pollen and spore terminology. LPP Contr. Ser., 1, Utrecht, 71 pp. Rao, A.N., Ong, E.T., 1974. Pollen morphology of certain tropical plants. J. Palynol. 10, 1^37. Reille, M., 1992. Pollen et Spores d’Europe et d’Afrique du Nord. Lab. Bot. Hist. Palynol., Marseille, 520 pp. Reille, M., 1995. Pollen et Spores d’Europe et d’Afrique du Nord. Suppl. 1. Lab. Bot. Hist. Palynol., Marseille, 327 pp. Reille, M., 1998. Pollen et Spores d’Europe et d’Afrique du Nord. Suppl. 2. Lab. Bot. Hist. Palynol., Marseille, 521 pp. Roubik, D.W., Moreno, J.E., 1991. Pollen and Spores of Barro Colorado Island. Monographs in Systematic Botany from the Missouri Botanical Garden 36, 1^270. Roure, J.M., 1985. Palinolog|¤a Ibe¤rica. Fam. 1 a 20. Cupressaceae a Betulaceae. Orsis 1, 43^69. Solereder, H., 1889. Aristolochiaceae. In: Engler, A., Prantl, K. (Eds.). Die natu«rlichen P£anzenfamilien, III Teil, 1. Abt. Verlag Wilhelm Engelmann, pp. 264^273. Stachurska, A., Sadowska, A., Kuszell, T., 1981. Kartoteka Palinologiczna Roslin Polskich. Opol. Tow. Przyj. Nauk Zesz. Przyr. 21, 257^266. Straka, H., 1981. Zur Pollenkunde (Palynologie) der in Band III/1 behandelten Familien. In: Hegi, G. (Ed.), Illustrierte Flora von Mittel-Europa. Band III. Teil 1, 3rd edn. (Conert, H.J., Hamann, U., Schultze-Motel, W., Wagenitz, G., Eds.). Verlag Paul Parey, Berlin, pp. 488^504. Tarnavschi, I.T., SVerba›nescu-Jitariu, G., Mitroiu-Ra›dulescu, N. and Ra›dulescu, D., 1981. A Monograph of Pollen in the Flora of Rumania. I (in Rumanian). Editura Academiei Roma›ne, Bucarest, 144 pp. Tutin, T.G., Burges, N.A., Chater, A.O., Edmondson, J.R., Heywood, V.H., Moore, D.M., Valentine, D.H., Walters,
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S.M., Webb, D.A. (Eds.), 1993. Flora Europaea, Volume 1: Psilotaceae to Platanaceae (2nd ed.). Cambridge University, pp. 87^89. Valde¤s, B., D|¤ez, M.J., Ferna¤ndez, I. 1987. Atlas polinico de Andalucia Occidental. Inst. Desarro. Reg. No. 43, Univ. Sevilla Excma. Dip. Cadiz, 450 pp. Walker, J.W., 1974. Aperture evolution in the pollen of primitive Angiosperms. Am. J. Bot. 61 (10), 1112^1137.
Walker, J.W., 1976. Evolutionary signi¢cance of the exine in the pollen of primitive angiosperms. In: Ferguson, I.K., Muller, J. (Eds.), The Evolutionary Signi¢cance of the Exine. Linnean Press, pp. 251^308. Wodehouse, R.P., 1935. Pollen Grains. McGraw-Hill, New York, 574 pp.
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Plate descriptions (all ¢gures U2000 except where otherwise stated) PLATE 1 (p. 52) Aristolochia clematitis L. (Campbell s.n.) 1. SEM micrograph; overall view. 2. SEM micrograph; ornamentation (U5000). Aristolochia paucinervis Pomel (Alston 10373) 3. SEM micrograph; overall view. 4. SEM micrograph; ornamentation (U5000). PLATE 2 (p. 53) Aristolochia paucinervis Pomel (¢g. 1, Alston 10373, ¢gs. 2^4, Van der Knaap and Van Leeuwen 89-4) 1. SEM micrograph; cross-section of the exine (U5000). 2. Cross-section. 3. Ornamentation at high focus. 4. Ornamentation at low focus. Aristolochia rotunda L. (Hekking 58-12A) 5. Cross-section. PLATE 3 (p. 54) Aristolochia rotunda L. (¢gs. 1^2, Davis and Sutton 64005; ¢gs. 3^4, Hekking 58-12A) 1. SEM micrograph; overall view. 2. SEM micrograph; ornamentation (U5000). 3. Ornamentation at high focus. 4. Ornamentation at low focus. PLATE 4 (p. 55) Asarum europaeum L. (¢gs. 1, Desplantes 19195; ¢gs. 2^5, Fresh Material Mulder Anno 1995) 1. SEM micrograph; polar view. 2. Polar view; ornamentation at high focus. 3. Polar view; ornamentation at medium focus. 4. Polar view; ornamentation at low focus. 5. Polar view, cross-section.
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PLATE 1 (Aristolochia clematitis type: Aristolochia clematitis, 1^2; Aristolochia paucinervis, 3^4)
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PLATE 2 (Aristolochia clematitis type: Aristolochia paucinervis, 1^4; Aristolochia rotunda, 5)
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PLATE 3 (Aristolochia clematitis type: Aristolochia rotunda)
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PLATE 4 (Asarum europaeum type: Asarum europaeum)
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