BS 55 423 Moraceae diversity in a global perspective Cornelis C. Berg Berg, C.C. 2005. Moraceae diversity in a global perspective. Biol. Skr. 55: 423-440. ISSN 0366-3612. ISBN 87-7304-304-4. The Moraceae are tightly embedded in the Urticales, a clear-cut entity of the Angiospermae. It comprises c. 1050 species and 37 genera, many of which have 1 or 2 species, and a few are large: Ficus has nearly 750 spp., Dorstenia has c. 105 spp., and Artocarpus has c. 60 spp. The family is pantropical, with extensions to subtropical and warm-temperate regions. It is well-represented in all tropical phytogeographic regions, each with one or more primary and/or secondary centres of distribution. The family is very diverse in both woody and herbaceous life and growth forms, but even more so in reproductive structures, comprising by condensation and fusion complex inflorescences, of which some are pseudoflorous and others are pseudocarpous, such as those of Ficus in which stigmas and anthers are not exposed at anthesis. This is linked to an unique polli­ nation system. For other Moraceae pollination is by wind and adapted to forest conditions, or they are pollinated by small beetles, flies, or thrips and based on breeding in staminate inflores­ cences, or the mode of pollination is unknown. Patterns of Moraceae diversification, its func­ tional implications, patterns of distribution, and the distinctness of genera and tribes suggest that most of the differentiation of the family took place in a still coherent tropical continent. The dis­ tinctness of taxa is not reflected in cladograms based on molecular analyses; one may wonder why. Cornelis C. Berg, The Norwegian Arboretum/Botanical Institute, University of Bergen, 5259 Hjellestad, Nor­ way. E-mail: [email protected]. - National Herbarium Nederland, Univ. Leiden branch, P.O. Box 9514, 2300 RA Leiden, The Netherlands. E-mail: [email protected] Introduction Urticales This paper summarizes various previously pub­ Except for doubt about the position of some lished aspects of the diversity and distribution odd genera or families (e.g, Barbeyaceae), the of Moraceae (Berg 1977a, 1983, 1989, 1990, Urticales was soon recognized and generally 1998, 2001) and discusses the overall taxo­ accepted as a well-defined group in the process nomic structure of the family and how they are of defining “natural” entities, as by Bentham linked to patterns of morphological and bio­ (1880), Bailion (1875), Eichler (1875), Engler logical diversification and distribution. Finally, (1889). The solidity of the order is in recent these morphological and biological aspects are times confirmed, e.g., by Baas et al. (2000) who compared with results of molecular analyses considered the wood anatomy as well as by (Sytsma et al. 2002). results of molecular analyses (Sytsma et al. 2002). 424 BS 55 The position of the order within the ply of food to frugivorous animals (Balslev & angiosperms has been uncertain. Traditionally Renner 1989; Gentry 1993; Berg 1998; Shana­ it is linked to the amentiferous families, han et al. 2001). grouped in the Hamamelidae (Cronquist 1981). A position near the Malvales (and Euphorbiaceae) has been advocated (Berg Composition of the family 1989). Molecular studies indicate a link to the The genera are very unequal in species num­ Rosales (c.g., Sytsma et al. 2002) and the order bers; 13 genera have only one or two closely is then referred to as Urticalean rosids. Roses related species; 14 genera have three to 14 and nettles in the same taxonomic unit are, species; few have more, Naucleopsis and Streblus although they share the capacity to sting, diffi­ 20-25 species, Artocarpus c. 60 species, Dorstenia cult to accept for morphological reasons. c. 105 species, and Ficus nearly 750 species and, therewith, one of the largest genera of woody flowering plants. Moraceae The Moraceae can be regarded as a “natural entity”, after exclusion of the genera in the Habit subfamilies Conocephaloideae and the The family is predominantly woody. The two Cannaboideae in Engler’s classification, the species of Fatoua are herbaceous. Eleven former being included in the Cecropiaceae species of Dorstenia are woody, the others are (Berg 1989) and the latter in the Cannabaceae herbaceous and represent a wide range of life (Cronquist 1981). and growth forms (Berg 1977a; Berg & Hijman In its morphological differentiation, the 1999; Fig. 2). family shows strong links to the other larger The woody Moraceae vary from being small urticalean families currently recognized. These shrubs to tall trees with sympodial or monopo­ links are such that one could unite Moraceae, dial growth. Few species are (sub) monocaul, Cecropiaceae, and Urticaceae into a single Dorstenia djettii], L. Guillaumet, Naucleopsis stip­ family. If one is not too narrow-minded, even ularis Ducke, and some Ficus species, e.g., F. Ulmaceae (both Celtidoideae and Ulmoideae) pseudopalma Blanco. Several Ficus species are can be included in this more broadly con­ rheophytic (van Steenis 1981). strued family Urticaceae, thus, largely coincid­ The capacity of Ficus to produce aerial ing with the order Urticales. adventitious roots, contributes considerably to However, such a broadly construed family the diversity of the woody life and growth lacks the transparency one needs to analyse forms, as hemi-epiphytes, hemi-epiliths, and morphological differentiation and distribution root-climbers. patterns in relation to biological traits and evo­ The plants have a system of ducts extending lutionary processes leading to its present taxo­ from the stem to the leaves and containing nomic structure. In this respect, it is conve­ milksap (not in Fatoua). Uncinate hairs occur nient to maintain a family Moraceae and its in many genera, and pluricellular (“glandu­ presumably natural subdivisions (Fig. 1). lar”) trichomes in most (or all?) genera. Nearly Compared to many other tropical plant fam­ all Ficus species have waxy glandular spots on ilies, the Moraceae is with its 1050-1100 species the lamina, base of the petiole and/or nodes of small, but in many tropical lowlands it is ecoli- leafy twigs. cally and important in forest structure and sup­ The height of the plants varies from less than BS 55 425 Fig. 1. Distribution and sub­ America Africa Asia Australasia division of Moraceae. The Moreae (ca 70) (11) (12) (c 47) survey gives the representa­ Madura (3*) Madura (1) Madura (7) tion in the three main phy­ Morus (3*) Morus (1) Morus (c 8) togeographic regions and Trophis (5) Trophis (2) Trophis (2) tribes with the numbers of Bleekrodea (1) Bleekrodea (1) species () per genus tribe, Broussonetia (1) Broussonetia (7) and region; in Ficus (Ficeae) Fatoua (1) Fatoua (1) per subgenus (for the Streblus (3) Streblus (21) Milicia (2) entirely or mainly Artocarpeae (c 70) (3) (c 68) (gyno) dioecious subgenera Treculia (3) (B) or for the (main) sec­ Artocarpus (c 60) tions of the monoecious Hullettia (2) subgenera (A) Ficus = Parartocarpus (2) Ficeae), and the indigenous Prainea (4) genera in [ ]; * including a Soroceae (23) (23) species in North America; o Bagassa (1) all (gyno) dioecious. Africa Batocarpus (3) includes Madagascar, adja­ Clarisia (3) cent Indian Ocean Islands, Poulsenia (2) and the Arabian Peninsula Sorocea (14) with adjacent islands. "Antiariopsidae" (3 (3) Antiaropsis (2) Sparattosyce ( 1 ) Dorstenieae (128) (64) (63) (1) Dorste nia (46) Dorstenia (58) Dorstenia (1) Bosqueiopsis ( 1 ) Scyphosyce (2) Trilepisium (1) Utsetela (1) Brosimum (14) Helianthostylis (2) Trymatococcus (2) Castilleae (58) (55) (2) (1) Antiaris (1) Antiaris (1) Mesogyne (1) Castilla (3) Helicostylis (7) Maquira (4) Naudeopsis (22) Perebea (9) Pseudolmedia (10) Ficeae (c.735) (c 120) (105) (c 510) "A" (c 350) (c 120) (83) (c 150) Pharmacosycea (c 20) Oreosycea (4) Oreosycea (c 50) Americana (c 100) Urostigma (7) Urostigma (c 80) Galoglychia (72) Stilpnophyllum (c 20) "B" (c 360) (22) (c 360) Ficus (1) Ficus (c 60) Synoecia (c 80) Sycidium (9) Sycidium (c 100) Sycomorus (12v) Sycomorus (c 120) 19[14] gen (c 270) 17[7]gen (185) 16[6] gen (c 610) 426 BS 55 WOODY PLANTS --------------------- > Herbaceous plants(1) TERRESTRIAL--- > Hemi-epiphytic(2) --- > Hemi-epilithic(2) TREES --------------------- > Shrubs (3) --- > --------------------- > Cli mb e r s ( 4 )-- ► Small Medium-sized (rarely with thorns or prickles) Emergents —► Climbers > Shrubs------ ► Rheophytic (5) Straggling(6) Obligatory With roots(7) Facultative With thorns(8) Erect -------- > Holo-epiphytic(9 Creeping (10) Herbaceous plants TERRESTRIAL------------------- ► Holo-epiphytic ( 11 ) PERRENIAL -------------------► Annual(12) NON-SUCCULENT ---------------- ► Succulent(13) Stem-succulent Tuber-succulent (geophytic or not) Phanerophytic (14) Geophytic(15) Erect Stem normal Not rosulate* Stem tuberous Rosulate* Creeping *Internodes long versus short Fig. 2. Life and habit forms of Moraceae. The numbers indicate that the life and habit forms are represented in the fol­ lowing genera and species: (1) Dorstenia p.maj.p. (2) Heus p.p. (3) Bleekrodea, Brosmium p.p., Ficus p.p., Helianthostylis p.p., Perebea p.p., Scyphosyce, Soroceap.p., Utsetlea. (4) Broussonetia kurzii, Ficus p.p., Madura p.maj.p., Prainea scandens, Trophis scan­ dens. (5) Ficus p.p. (6) Broussonetia kurzii, Ficus p.p., Prainea scandens, Trophis scandens. (7) Ficus p.p. (8) Madura p.p. (9) Ficus deltoidea (and F. oleifolid), facultatively. (10) Ficus p.p. (11) Dorstenia astyanactis. (12) Dorstenia annua. (13) Dorstenia p.p., Fatoua. (14) Dorstenia p.p. 5