THE GLOBAL FLORA A practical flora to vascular plant species of the world

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3 Plant Gateway s THE GLOBAL FLORA ANGIOSPERMS 232. PETENAEACEAE by J.W. BYNG & M.J.M. CHRISTENHUSZ Field images by Christopher Davidson January 2018

4 Angiosperms, Petenaeaceae Vol. 2: Published by Plant Gateway Ltd., 5 Baddeley Gardens, Bradford, BD10 8JL, United Kingdom Plant Gateway 2018 This work is in copyright. Subject to statutory exception and to the provision of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Plant Gateway Ltd. ISSN eissn Plant Gateway has no responsibility for the persistence or accuracy of URLS for external or third-party internet websites referred to in this work, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate. British Library Cataloguing in Publication data A Catalogue record of this book is available from the British Library For information or to purchase other Plant Gateway titles please visit Authors James W. Byng, Plant Gateway, Bradford & Kingston, United Kingdom and Den Haag, the Netherlands; Naturalis Biodiversity Center, Leiden, The Netherlands. Maarten J.M. Christenhusz, Plant Gateway, Bradford & Kingston, United Kingdom and Den Haag, the Netherlands; Royal Botanic Gardens, Kew, United Kingdom. Cover image: Petenaea cordata Christopher Davidson

5 Summary Petenaeaceae is endemic to Central America and contains one species in one genus. The sole species, Petenaea cordata, was originally described in Elaeocarpaceae, but was also previously treated in Tiliaceae (= Malvaceae sensu APG IV) or associated with Muntingiaceae, all based on superficial resemblances. A recent molecular study found the genus to belong to Huerteales, sister to African Gerrardinaceae, with which it does not share any significant synapomorphies. Diagnostic macromorphological characters for the family are: pubescent trees or shrubs with simple, alternate leaves, palmatinerved and with cordate leaf bases and minute stipules; flowers with pink sepals, petals absent, but the receptacle is adorned with long, showy, bright pink, multicellular hairs, and superior ovaries; and lobed edible berries. The wood vessels are in radial multiples, fibres are conspicuously septate, and rays are uni- to multiseriate and of two distinct sizes. Pollen grains, described here for the first time, are spheroidal, tricolporate and with perforate ornamentation. The sole species in the family is illustrated and comes with a description including data on its habitat, known herbarium specimens at major herbaria, distribution map and additional observations. 1

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7 PETENAEACEAE Christenh., M.F.Fay & M.W.Chase (in Christenhusz et al. 2010: 23) Large shrubs or trees, hairy; trichomes multicellular, simple or branched, white. Stems with vessels in radial multiples, fibres conspicuously septate. Leaves simple, alternate, cordate, margins minutely denticulate, palmatinerved, long-petiolate, petioles tinged magenta; stipules minute, triangular. Inflorescences many-flowered, villous-tomentose, paniculate cymes; peduncles and pedicels tinged magenta, covered in white hairs. Flowers bisexual, actinomorphic. Sepals (4 or) 5, valvate, pink and white hairy outside. Petals absent. Receptacle covered in a fringe of long multicellular, bright pink, showy hairs. Stamens 8 12, free, borne on a fleshy annular disk; nectary glands placed between the glabrous filaments; anthers dorsifixed, dehiscing by apical slits. Pollen spheroidal, tricolporate. Ovary superior, carpels 4-5, fused, placentation axile, ovules numerous; style terminal topped with a disc-shaped stigma. Fruit an edible berry, more or less globose, somewhat lobed, ripening magenta to maroon. Seeds minute, numerous. The family contains one genus and one species: Petenaea cordata Lundell Distribution Endemic to Central America, where it occurs in western Belize, northern Guatemala and southeastern Mexico (Chiapas, Tabasco) (Figure 1). Figure 1: Global distribution of Petenaeaceae (highlighted). 3

8 Classification The genus was originally described in the family Elaeocarpaceae based on the baccate fruits, and comparisons of wood anatomy, but it was mentioned that it was similar to Muntingia L., which was then also placed there. Later Petenaea was placed in Tiliaceae (now Malvaceae), probably because of superficial resemblance to Tilia L.. It was always difficult to place on morphological grounds due to its unusual combination of characters: simple and branched trichomes, palmately veined leaves with minute stipules, petals absent, but a crown of moniliform hairs replace it, nectary glands between the stamens, dorsifixed anthers and a large axile placenta with numerous ovules forming a berry-like fruit. Muntingiaceae was the closest morphological fit, which is why it was placed there tentatively, but the lack of petals and differences in wood anatomy made this relationship unlikely. Molecular analysis indicate it to be a member of Huerteales (Figure 2), a recent order that includes a number of family misfits. Its closest relative is Gerrardina Oliv. (Gerrardinaceae) a former Flacourtiaceae from Africa, with which it shares the leaf shape and berry-like fruit, but few other morphological characters support this relationship. Petenaea cordata has no close relatives and is probably a relic of a formerly more diverse lineage. Wood anatomy (Figure 3) The heartwood is pale reddish-brown and more or less sharply demarcated from the sapwood. Growth ring boundaries are absent or indistinct. The wood is diffuse-porous with vessels mostly in radial multiples of 2 ( 12). The vessel perforation plates are simple and oblique; intervessel pits are alternate and about 8 μm in horizontal diameter; vessel-ray have reduced borders and are often scalariform. Tangential diameter of vessel lumina is μm and mean vessel element length is 635 μm. Tyloses present in the heartwood. The fibres are thick-walled, conspicuously septate, and with small minutely bordered pits. Mean fibre length is 1130 μm. Axial parenchyma is rare or absent. The rays are both uni- and multiseriate, 1 5 cells in width and up to 40 cells high, strongly heterocellular with many square and upright cells but body cells are often procumbent. Crystals and minerals are absent. 4

9 Angiosperms Eudicots Monocots Rosids Superrosids Asterids Superasterids Amborellales Nymphaeales Austrobaileyales Cannellales Piperales Magnoliales Laurales Chloranthales Acorales Alismatales Petrosaviales Pandanales Dioscoreales Liliales Asparagales Arecales Commelinales Zingiberales Poales Ceraotophyllales Ranunculales Proteales Trochodendrales Buxales Gunnerales Dilleniales Saxifragales Vitales Zygophyllales Fabales Rosales Fagales Cucurbitales Celastrales Oxalidales Malpighiales Geraniales Myrtales Crossosomatales Picramniales Sapindales Huerteales Malvales Brassicales Berberidopsidales Santalales Caryophyllales Cornales Ericales Icacinales Metteniusales Garryales Gentianales Boraginales Vahliales Solanales Lamiales Aquifoliales Asterales Escalloniales Bruniales Paracryphiales Dipsacales Apiales 231. Gerrardinaceae 232. Peteneaceae Petenaea 233. Tapisciaceae 234. Dipentodontaceae Figure 2: Classification of Petenaeaceae showing the order and generic rank relevant to the family. Adapted from APG IV (2016) and Christenhusz et al. (2010). 5

10 A B 60 μm 80 μm C 100 μm Figure 3: Wood anatomy of Petenaea cordata. A, transverse section showing vessels arranged mainly in radial multiples; B, tangential longitudinal section showing a mix of uni- and ultiseriate rays; C, radial longitudinal section showing vessel-ray pitting and septa of the fibres. Images taken from Kurachka (1962) (CC BY-NC-SA 4.0). 6

11 Pollen morphology The pollen of Petenaea (Figure 4) is shed as small-sized monads (polar axis [P] by equatorial axis [E] = by μm). Pollen grain shape is spheroidal in hydrated condition and prolate with lobate outline in dry condition. The aperture system is tricolpate, apertures sunken in dry condition. The ornamentation is perforate with minute pits. Methods: Pollen grains from herbarium material were mounted on stubs using double-sided adhesive tape, sputter-coated with platinum and observed with the SEM. Specimen examined: Pennington & McQueen The pollen of Petenaea is, as far as we aware, described here for the first time. Pollen grains of Petenaea are similiar to others species in Huerteales, shedding as monads, small in size (10 25 μm) and with tricolporate apertures, but the exine structure is either more or less perforate (Tapiscia Oliv.) or reticulate (Gerrardina, Huertea Ruiz & Pavon) (Keating, 1973; Dickison, 1987). A B C 2 μm 2 μm 2 μm Figure 4: Dry pollen grains of Petenaea cordata. A, polar view; B & C, equatorial views. Images original. References APG IV (2016), Bayer (2003), Christenhusz (2010), Christenhusz et al. (2010), Dickison (1987), Keating (1973), Kukachka (1962), Lundell (1962). 7

12 1. Petenaea Lundell (1962: 22) Type Petenaea cordata Lundell Etymology Petenaea is named after the Guatemalan department El Petén, from where the genus was first described. Description as for the family. 1. Petenaea cordata Lundell (1962: 24) Figures 5, 6, 7. Table 1. Type GUATEMALA. Petén, Lake Petén Itza, along northern shore, between San José and El Remate, 23 January 1962, C.L. Lundell (holotype LL [LL ]; isotypes CAS [CAS ], F [F F], G [G ], K [K ], LL [LL ], MEXU [MEXU ], S [S ], US [US ]). Etymology cordata, referring to the heart-shaped leaves. Trees to ca. 10 m or large shrubs. Stems often tinged red, villous-tomentose. Leaves minutely stipulate; petioles 5 11 cm, densely short villous, magenta; blades cm, chartaceous, densely villous below, glabrescent above, turning red with age, palmately veined at base with 5 7 primary veins, the secondary venation reticulate, the base broadly cordate, the margins minutely denticulate, the apices acute to broadly short-acuminate. Stipules minute, soon caducous. Inflorescences axillary, cymose, up to 6 cm, longpedunculate, rose-pink, the branches villous-tomentose; pedicels 5-12 mm, pink. Flowers with an annular disk, glandular; buds up to 4 mm, lanceolateattenuate. Sepals 4 or 5, ca. 4 mm, valvate, lanceolate, attenuate from the base to the apex, reflexed, magenta-pink, the base bearing 2 3 obovoid subsessile glands, and densely villous with c. 2 mm long, moniliform, hotpink hairs. Petals absent. Stamens 8 12, mm, glabrous; anthers ca. 1.5 mm, dorsifixed, lanceolate-oblong, introrse, yellow, opening by an apical pore-like slit. Ovary sessile, tomentose; style slender, pubescent at base; 8

13 A B C D E F Figure 5: Living images of Petenaea cordata. A, habit; B, lower leaf surface; C, close-up of flower and buds; D, inflorescence; E, mature and immature fruits; F, cross-section of fruits. All images Christopher Davidson. 9

14 A B 1 cm 1 cm D 1 cm C 5 mm Figure 6: Herbarium images of Petenaea cordata. A, Upper leaf surface; B, inflorescence; C, close-up of flowers; D, lower leaf surface. Images: A, B & D - Lundell 17272, C - Pennington & McQueen Images original. 10

15 stigma discoid. Berry 6 12 mm, ovoid to subglobose, shallowly 4 5 lobed, pulpy, sweet, sparsely pubescent, turning deep red to maroon-black when mature; style persistent. Seeds numerous, c. 1 mm long, oblong-pyramidal or irregular. Distribution 79: MXT; 80: BLZ GUA. The species is known from southeast Mexico to northern Guatemala and western Belize. It is most abundant in Chiapas. Habitat and ecology Found in forests bordering lakeshores, streams and rivers, on gypsum escarpments, and in open, disturbed secondary forests and along roadsides; m elevation. Phenology Flowering and fruiting continuously throughout the year. Fruits are edible and are locally consumed by people. Chromosome number and genome size unknown. Preliminary conservation status Least concern (LC) according to the IUCN Red List Criteria (IUCN, 2014), as the species is locally abundant and faces no immediate threats. References Christenhusz (2010), Christenhusz et al. (2010), IUCN (2014), Lundell (1962). Figure 7: Distribution of Petenaea cordata in Central America based on herbarium records listed in Table 1. 11

16 Table 1: Herbarium specimens consulted at BM, K, L, MO, P and U of Petenaea cordata are listed and arranged by geography followed by collector and collectors number. Duplicates and specimens at other herbaria are listed where known. Country Department Municipality Collector Collector number Collection date Herbarium Elevation (m) Phenology Belize Cayo Benque Viejo Contreras, E Oct-1967 MO Fl Belize Cayo Ceibo Creek Peña-Chocarro, M. et al Mar-2000 BM, BRH, MEXU, MO, PMA 879 Fl Belize Toledo Bladen Brewer, S.W Apr-2010 MO 60 Fl & Fr Belize Toledo Bladen Brewer, S.W. & Rejmánek, M May-2002 MO 350 Fl & Fr Guatemala Alta Verapaz Chisec Morales Can, J. et al Feb-2005 MO, USCG 408 Guatemala Huehuetenango Nenton Christenhusz, M.J.M. et al Mar-2009 BM, H, L, MO, USCG Fl Guatemala Petén Dolores Tún Ortíz, R Feb-1971 MO Immature Fr Guatemala Petén Flores Peña-Chocarro, M. et al Feb-2000 BM, MO Fl & Fr Guatemala Petén Flores Pennington, T. & McQueen Feb-1991 K, U 220 Fl & Fr Guatemala Petén Flores Tún Ortíz, R Feb-1970 MO Fl Guatemala Petén Flores Tún Ortíz, R Feb-1970 P Fl Guatemala Petén Poptún Lundell, C.L. & Contreras, E Sep-1976 MO Fr Guatemala Petén Poptún Contreras, E Mar-1970 MO Fl & Fr Guatemala Petén San Jose Contreras, E Jan-1962 P Fl Guatemala Petén San Jose Contreras, E Aug-1966 MO, U Fl Guatemala Petén San Jose Contreras, E Aug-1967 U Fl Guatemala Petén San Jose Lundell, C.L Jan-1962 BM, G, P Fl & Fr Guatemala Petén San Jose Lundell, C.L Jan-1962 BM Guatemala Petén San Jose Lundell, C.L Jan-1962 K, LL, MO Fl Guatemala Petén San Luis Contreras, E Feb-1971 MO Fl & immature Fr 12

17 Table 1 continued Country Department Municipality Collector Collector number Collection date Herbarium Elevation (m) Phenology Guatemala Petén San Luis Contreras, E Feb-1971 MO Fr Mexico Chiapas Chilón Ishiki, M. et al Nov-1998 U 850 Fl & immature Fr Mexico Chiapas Chilón Téllez V., O. et al May-1982 MEXU, MO Fr Mexico Chiapas Ocosingo Aguilar M., G May-2002 MO 325 Fl & Fr Mexico Chiapas Ocosingo Aguilar M., G. & Álvarez M., D Oct-2002 MO 330 Mexico Chiapas Ocosingo Aguilar M., G. & Chancayun, C Apr-2003 MO 210 Immature Fr Mexico Chiapas Ocosingo Aguilar M., G. et al Mar-2002 MO Fl & immature Fr Mexico Chiapas Ocosingo Aguilar M., G. et al Jan-2003 MO 305 Fl & Fr Mexico Chiapas Ocosingo Aguilar M., G. et al Aug-2002 MO 325 Fl Mexico Chiapas Ocosingo Aguilar M., G. et al Sep-2002 MEXU, MO 220 Fl & immature Fr Mexico Chiapas Ocosingo Aguilar M., G. et al Oct-2002 MO 383 Fl & immature Fr Mexico Chiapas Ocosingo Aguilar M., G. et al Dec-2002 MEXU, MO 386 Fl & immature Fr Mexico Chiapas Ocosingo Aguilar M., G. et al Sep-2003 MO 431 Fl & immature Fr Mexico Chiapas Ocosingo Alush Shilom Ton Feb-1983 MO 600 Fl & immature Fr Mexico Chiapas Ocosingo Álvarez M., D. & Aguilar M., G Aug-2003 MO 228 Fl Mexico Chiapas Ocosingo Álvarez M., D. & Chambor, A Apr-2003 MO 210 Fr Mexico Chiapas Ocosingo Álvarez, D. et al Nov-2003 MEXU, MO 257 Fl & immature Fr Mexico Chiapas Ocosingo Calónico Soto, J Oct-2002 MEXU, MO 371 Fl & Fr Mexico Chiapas Ocosingo Calónico Soto, J. et al Nov-2002 MEXU, MO 190 Fl Mexico Chiapas Ocosingo Hammel, B.E Oct-1986 MO 780 Fl & Fr Mexico Chiapas Ocosingo Martínez S., E.M Aug-1984 MO 220 Fl & immature Fr Mexico Chiapas Ocosingo Martínez S., E.M Dec-1984 MEXU, MO 80 13

18 Table 1 continued Country Department Municipality Collector Collector number Collection date Herbarium Elevation (m) Phenology Mexico Chiapas Ocosingo Martínez S., E.M Dec-1984 MO 120 Fl & immature Fr Mexico Chiapas Ocosingo Martínez S., E.M Feb-1985 MO 180 Fl Mexico Chiapas Ocosingo Martínez S., E.M Apr-1985 MEXU, MO 180 Fl bud Mexico Chiapas Ocosingo Martínez S., E.M Jun-1986 MO 370 Immature Fr Mexico Chiapas Ocosingo Martínez S., E.M Feb-1986 MO 980 Fl Mexico Chiapas Ocosingo Martínez S., E.M Apr-1986 MEXU, MO 700 Fr Mexico Chiapas Ocosingo Torres C., R. & Cedillo T., R Dec-1982 MO 540 Fl & immature Fr Mexico Chiapas Palenque Alush Shilom Ton Nov-1982 MO Fl & immature Fr Mexico Chiapas Palenque Breedlove, D.E. & Strother, J.L Oct-1980 MO 300 Fl & Fr Mexico Chiapas Palenque Mexico Chiapas Palenque Cabrera C., E.F. & Cabrera, H. de Cabrera C., E.F. & Cabrera, H. de Aug-1983 MEXU, MO Fl Aug-1984 MEXU, MO 364 Mexico Chiapas Palenque Cabrera C., E.F. et al Mar-1982 MEXU, MO Fl bud Mexico Chiapas Palenque Cabrera C., E.F. et al May-1982 MEXU, MO 211 Mexico Chiapas Palenque Charles H. Perino & Janice V. Perino Jun-1978 MO 400 Fl & Fr Mexico Chiapas Palenque Chehaibar N., T Aug-1985 MO 250 Fl Mexico Chiapas Tumbalá Stevens, W.D. & Martínez S., E.M Sep-1988 BM, MO 810 Fr Mexico Chiapas Tumbalá Téllez V., O. & Martínez S., E.M Sep-1979 MEXU, MO Mexico Chiapas Yajalón Cabrera C., E.F. & Cabrera, H. de Dec-1983 MEXU, MO 544 Fl & Fr Mexico Chiapas Yajalón Cabrera C., E.F. & Huft, M.J Oct-1984 MEXU, MO 650 Fl & Fr Mexico Tabasco Tacotalpa Cowan, C.P. et al Mar-1980 MO 300 Fl & immature Fr Mexico Tabasco Tenosique Cowan, C.P. & R. Curiel, R May-1984 MO 270 Fr 14

19 REFERENCES APG IV. (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: Bayer, C. (2003) Muntingiaceae. In: Kubitzki, K. (ed.). Families and Genera of Vascular Plants 5: Christenhusz, M.J.M. (2010) Petenaeaceae. Draft manuscript for Flora Mesoamericana 3 (2). Published online pdf?projectid=3&langid=66 Christenhusz, M.J.M., Fay, M.F., Clarkson, J.J., Gasson, P., Morales Can, J., Jiménez Barrios, J.B. & Chase, M.W. (2010) Petenaeaceae, a new angiosperm family in Huerteales with a distant relationship to Gerrardina (Gerrardinaceae). Botanical Journal of the Linnean Society 164: Dickison, W.C. (1987) A palynological study of the Staphyleaceae. Grana 26: IUCN. (2014) IUCN Red List Categories and Criteria: Version nd Edition. IUCN Species Survival Commission, IUCN Gland & Cambridge. Keating, R.C. (1973) Pollen morphology and relationships of the Flacourtiaceae. Annals of Missouri Botanic Gardens 60: Kukachka, B.F Wood anatomy of Petenaea cordata Lundell (Elaeocarpaceae). Wrightia 3: Lundell, C.L. (1962) Plantae Mayanae V. Petenaea cordata, a new genus and species in the Elaeocarpaceae, and other taxonomic notes. Wrightia 3: acknowledgements We thank the curators of BM, K, L, P and U for access to their collections. We also thank Chris Davidson for allowing us to use his field photos. Lastly, we thank Frederic Lens for his advice on wood anatomy. 15

20 INDEX TO SCIENTIFIC NAMES Elaeocarpaceae.... 1, 4 Flacourtiaceae Gerrardina Oliv , 7 Gerrardinaceae... 1, 4 Huerteales... 1, 4, 7 Huertea Ruiz & Pavon Malvaceae... 1, 4 Muntingia L Muntingiaceae... 1, 4 Petenaea cordata Lundell...1, 3, 4, 6, 7, 8, 9, 10, 11, 12 Petenaea Lundell...1, 4, 7, 8 Petenaeaceae... 1, 3, 5 Tapiscia Oliv Tilia L Tiliaceae.... 1, 4 16

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22 THE GLOBAL FLORA Series 1 Lycopods Isoëtaceae FERNS Psilotaceae Pteridaceae Saccolomataceae Salviniaceae Schizaeaceae Lycopodiaceae Selaginellaceae Aspleniaceae Cyatheaceae Cystodiaceae Dennstaedtiaceae Dipteridaceae Equisetaceae Gleicheniaceae Hymenophyllaceae Lindsaeaceae Lonchitidaceae Marattiaceae Marsileaceae Matoniaceae Ophioglossaceae Osmundaceae Polypodiaceae GYMNOSPERMS Araucariaceae Cupressaceae Cycadaceae Ephedraceae Ginkgoaceae Gnetaceae Pinaceae Podocarpaceae Sciadopityaceae Taxaceae Welwitschiaceae Zamiaceae Acanthaceae Achariaceae Achatocarpaceae Acoraceae Actinidiaceae Adoxaceae Aextoxicaceae Aizoaceae Akaniaceae Alismataceae Alseuosmiaceae Alstroemeriaceae Altingiaceae Alzateaceae Amaranthaceae Amaryllidaceae Amborellaceae Anacampserotaceae Anacardiaceae Ancistrocladaceae Anisophylleaceae Annonaceae Aphanopetalaceae Aphloiaceae Apiaceae Apocynaceae Apodanthaceae Aponogetonaceae Aquifoliaceae Araceae Araliaceae Arecaceae Argophyllaceae Aristolochiaceae Asparagaceae Asphodelaceae Asteliaceae Asteraceae Asteropeiaceae Atherospermataceae Austrobaileyaceae Balanopaceae Balanophoraceae Balsaminaceae Barbeuiaceae Barbeyaceae Basellaceae Bataceae Begoniaceae Berberidaceae Berberidopsidaceae Betulaceae Biebersteiniaceae Bignoniaceae Bixaceae Blandfordiaceae Bonnetiaceae Boraginaceae Boryaceae Brassicaceae Bromeliaceae Brunelliaceae Bruniaceae Burmanniaceae Burseraceae Butomaceae Buxaceae Byblidaceae Cabombaceae Cactaceae Calceolariaceae Calophyllaceae Calycanthaceae Calyceraceae Campanulaceae Campynemataceae Canellaceae Cannabaceae Cannaceae Capparaceae Caprifoliaceae Cardiopteridaceae Caricaceae Carlemanniaceae Caryocaraceae Caryophyllaceae Casuarinaceae Celastraceae Centroplacaceae Cephalotaceae Ceratophyllaceae Cercidiphyllaceae Chloranthaceae Chrysobalanaceae Circaeasteraceae Cistaceae Cleomaceae Clethraceae Clusiaceae Colchicaceae Columelliaceae Combretaceae Commelinaceae Connaraceae Convolvulaceae Coriariaceae Cornaceae Corsiaceae Corynocarpaceae Costaceae Crassulaceae Crossosomataceae Crypteroniaceae Ctenolophonaceae Cucurbitaceae Cunoniaceae Curtisiaceae Cyclanthaceae Cymodoceaceae Cynomoriaceae Cyperaceae Cyrillaceae Cytinaceae Daphniphyllaceae Dasypogonaceae Datiscaceae Degeneriaceae Diapensiaceae Dichapetalaceae Didiereaceae Dilleniaceae Dioncophyllaceae Dioscoreaceae Dipentodontaceae Dipterocarpaceae Dirachmaceae Series 2 ANGIOSPERMS

23 Doryanthaceae Droseraceae Drosophyllaceae Ebenaceae Ecdeiocoleaceae Elaeagnaceae Elaeocarpaceae Elatinaceae Emblingiaceae Ericaceae Eriocaulaceae Erythroxylaceae Escalloniaceae Eucommiaceae Euphorbiaceae Euphroniaceae Eupomatiaceae Eupteleaceae Fabaceae Fagaceae Flagellariaceae Fouquieriaceae Francoaceae Frankeniaceae Garryaceae Geissolomataceae Gelsemiaceae Gentianaceae Geraniaceae Gerrardinaceae Gesneriaceae Gisekiaceae Gomortegaceae Goodeniaceae Goupiaceae Griseliniaceae Grossulariaceae Grubbiaceae Guamatelaceae Gunneraceae Gyrostemonaceae Haemodoraceae Halophytaceae Haloragaceae Hamamelidaceae Hanguanaceae Heliconiaceae Helwingiaceae Hernandiaceae Himantandraceae Huaceae Humiriaceae Hydatellaceae Hydrangeaceae Hydrocharitaceae Hydroleaceae Hydrostachyaceae Hypericaceae Hypoxidaceae Icacinaceae Iridaceae Irvingiaceae Iteaceae Ixioliriaceae Ixonanthaceae Joinvilleaceae Juglandaceae Juncaceae Juncaginaceae Kewaceae Kirkiaceae Koeberliniaceae Krameriaceae Lacistemataceae Lamiaceae Lanariaceae Lardizabalaceae Lauraceae Lecythidaceae Lentibulariaceae Lepidobotryaceae Liliaceae Limeaceae Limnanthaceae Linaceae Linderniaceae Loasaceae Loganiaceae Lophiocarpaceae Lophopyxidaceae Loranthaceae Lowiaceae Lythraceae Macarthuriaceae Magnoliaceae Malpighiaceae Malvaceae Marantaceae Marcgraviaceae Martyniaceae Maundiaceae Mayacaceae Mazaceae Melanthiaceae Melastomataceae Meliaceae Menispermaceae Menyanthaceae Metteniusaceae Microteaceae Misodendraceae Mitrastemonaceae Molluginaceae Monimiaceae Montiaceae Montiniaceae Moraceae Moringaceae Muntingiaceae Musaceae Myodocarpaceae Myricaceae Myristicaceae Myrothamnaceae Myrtaceae Nartheciaceae Nelumbonaceae Nepenthaceae Neuradaceae Nitrariaceae Nothofagaceae Nyctaginaceae Nymphaeaceae Nyssaceae Ochnaceae Olacaceae Oleaceae Onagraceae Oncothecaceae Opiliaceae Orchidaceae Orobanchaceae Oxalidaceae Paeoniaceae Pandaceae Pandanaceae Papaveraceae Paracryphiaceae Passifloraceae Paulowniaceae Pedaliaceae Penaeaceae Pennantiaceae Pentadiplandraceae Pentaphragmataceae Pentaphylacaceae Penthoraceae Peraceae Peridiscaceae Petenaeaceae 2: 1-16 Petermanniaceae Petrosaviaceae Phellinaceae Philesiaceae Philydraceae Phrymaceae Phyllanthaceae Phyllonomaceae Physenaceae Phytolaccaceae Picramniaceae Picrodendraceae Piperaceae Pittosporaceae Plantaginaceae Platanaceae Plocospermataceae Plumbaginaceae Poaceae Podostemaceae Polemoniaceae Polygalaceae Polygonaceae Pontederiaceae Portulacaceae Posidoniaceae Potamogetonaceae Primulaceae Proteaceae Putranjivaceae Quillajaceae Rafflesiaceae Ranunculaceae Rapateaceae Resedaceae Restionaceae Rhabdodendraceae Rhamnaceae Rhizophoraceae Ripogonaceae Rivinaceae Roridulaceae Rosaceae Rousseaceae Rubiaceae Ruppiaceae Rutaceae Sabiaceae Salicaceae Salvadoraceae Santalaceae Sapindaceae Sapotaceae Sarcobataceae Sarcolaenaceae Sarraceniaceae Saururaceae Saxifragaceae Scheuchzeriaceae Schisandraceae Schlegeliaceae Schoepfiaceae Scrophulariaceae Setchellanthaceae Simaroubaceae Simmondsiaceae Siparunaceae Sladeniaceae

24 Smilacaceae Solanaceae Sphaerosepalaceae Sphenocleaceae Stachyuraceae Staphyleaceae Stegnospermataceae Stemonaceae Stemonuraceae Stilbaceae Strasburgeriaceae Strelitziaceae Stylidiaceae Styracaceae Surianaceae Symplocaceae Talinaceae Tamaricaceae Tapisciaceae Tecophilaeaceae Tetracarpaeaceae Tetrachondraceae Tetramelaceae Tetrameristaceae Theaceae Thomandersiaceae Thurniaceae Thymelaeaceae Ticodendraceae Tofieldiaceae Torricelliaceae Tovariaceae Trigoniaceae Trimeniaceae Triuridaceae Trochodendraceae Tropaeolaceae Typhaceae Ulmaceae Urticaceae Vahliaceae Velloziaceae Verbenaceae Violaceae Vitaceae Vochysiaceae Winteraceae Xeronemataceae Xyridaceae Zingiberaceae Zosteraceae Zygophyllaceae Series 3 SPECIAL EDITIONS Introduction 1: 1-35

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