Moraceae

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Moraceae

Description

— The family consists chiefly of lowland trees. In Malesia, tall canopy trees or emergents are found in Antiaris, Artocarpus, Ficus, Parartocarpus, Prainea, and Streblus. They often have buttresses. Smaller, undergrowth trees of varying stature belong to Antiaropsis, Broussonetia, Ficus, Hullettia, Streblus, and Trophis. Shrubs and treelets are found in Bleekrodea, Ficus, Maclura, and Streblus. Species which can become tall trees, such as Streblus elongatus, may flower as shrubs or treelets.

Most tree species are evergreen, but some are deciduous both in seasonal and ever- wet forest. The trees have intermittent growth, with sometimes conspicuous buds that are temporarily dormant.

Monocaulous or sparingly branched trees, with relatively thick branches and terminal tufts of large leaves (pachycladous, or in Corner’s terminology pachycaul, trees), occur in several species of Ficus (e.g., F. pseudopalma), one species of Dorstenia (Africa), and one species of Naucleopsis (America). A wide range of tree forms occur between the pachycladous tree and the more common leptocladous one, with slender branches and small leaves mostly evenly distributed on the leafy twigs.

There are c. 100 species of climbers in Malesia, many of them belonging to Ficus and Maclura. Moreover, this habit is found in single species in Broussonetia and Trophis, and sometimes in Prainea scandens. The climbing habit is rare in Africa and America, in each restricted to a single species of Maclura and to one or two (sub)lianescent Ficus species. The climbers may be twining or straggling (Trophis scandens and species of Ficus subg. Sycidium sect. Palaeomorphe), scramblers with thorns (Maclura), or root climbers (Ficus subg. Synoecia).

Most species are unarmed, although thorns are found in Maclura and Streblus, in the former often to assist climbing. Spiny leaf margins and/or apices occur in some species of Streblus and also in the neotropical genera Clarisia and Sorocea.

Fatoua is predominantly herbaceous as are the majority of Dorstenia. In the latter genus there are a wide range of herbaceous life forms: stem succulents, tuber succulents, geophytes, epiphytes, and annuals (Berg & Hijman 1999). In both of these genera, plants can be suffrutescent. Suffrutescence is also known in the essentially woody Ficus (e.g., F. griffithii Miq. (Asia) and F. suffruticosa Corner (New Guinea)) and the neotropical Perebea (P. humilis C.C. Berg).

Hemi-epiphytism is a prominent life form in Ficus, which also has two species that can be holo-epiphytes. The very wide range of life and growth forms in Ficus are described under the genus.

The twig apices are shed in Morus and Broussonetia, and elongation of twigs is performed by the meristem in the upper most lateral bud; the resting buds are scaled. This trait is also found in other elements of northern temperate forest, such as Tilia and Ulmus. Shoot apices are even abscised in tropical lowland species of the two moraceous genera.
Trees, shrubs, woody climbers, or herbs (Dorstenia, Fatoua), terrestrial, hemi-epiphytic (or holo-epiphytic), dioecious or monoecious, with milky sap. Leaves spirally arranged or distichous, (sub)opposite or subverticillate; — The leaves are mostly alternate and spirally arranged or distichous. Opposite leaves are found in some groups of Ficus (subg. Sycidium and subg. Sycomorus), often in combination with alternate arrangement. Opposite leaves may also occur in Broussonetia papyrifera and are characteristic for the neotropical genus Bagassa. Subverticillate leaves occur in some species of Ficus subg. Sycidium. Such arrangement of the leaves occurs occasionally in Artocarpus (subg. Pseudojaca).

The lamina varies from large, up to 2 m long in Ficus solomonensis or up to 1 m wide in F. dammaropsis, to very small, not more than 3 cm long in F. vaccinioides King from Taiwan, and from 2 cm in F. humbertii C.C. Berg from Madagascar, or some neotropical Dorstenia species. The lamina is usually basally attached (peltate in some species of Dorstenia). It is mostly entire, less commonly pinnately or palmately incised. Pinnately incised laminas occur in some species of Artocarpus, Dorstenia, and Ficus, palmately incised ones in Broussonetia, Dorstenia, Ficus, and Morus. The venation is mostly brochidodromous. The tertiary venation is basically scalariform, with numerous parallel transverse veins in the intercostal area. This type of venation evolved into a reticulate pattern, and, in some groups of Ficus, subsequently into tertiary venation largely parallel to the secondary venation.

The texture of the lamina varies from thickly coriaceous to chartaceous, or even membranaceous in herbaceous genera. The margin is usually entire in coriaceous laminas (of evergreen forest species) and often dentate in thinner ones. The petiole varies from long to short. Long petioles are generally associated with cordiform laminas.

The stipules are often fully amplexicaul, leaving annular scars, by which so many Moraceae can be recognized, but they may also be semi-amplexicaul or lateral. The stipules vary from large (more than 10 cm long) to small (down to 1 mm long). They are mostly caducous but may be persistent or subpersistent.
stipules fully amplexicaul or semi-amplexicaul and lateral or intrapetiolar, free or connate; — The inflorescences are mostly paired as in other families of the Urticales. The types of inflorescences range from cymes, to racemes, to spikes (slender or thick and almost spadix-like), to heads, either globose- to ellipsoid-capitate, or discoid (bisexual or unisexual, involucrate or not), or to urceolate structures (opening or remaining closed).

They have a simple structure: racemose, spicate, cymose, or capitate, and mostly unisexual in the tribe Moreae and in the neotropical genera of the tribe Artocarpeae. The staminate ones often resemble catkins of Amentiferae, but the pistillate ones in fruit look different owing to the fleshy pericarp, perianths and/or interfloral bracts. In the majority of the Moraceae the inflorescences are complicated by condensation of axes, by fusion of homologous or adjacent structures, more prominent bracts, and bisexuality of inflorescences. These complex structures are mostly the functional entities with regard to pollination, and in fruiting state, to dispersal, as is very clearly the case in the syconium or fig of Ficus. The complex inflorescences can be pseudoflorous, but are more often pseudocarpous and they usually bear numerous flowers. Reduction to the uniflorous state sometimes occurs (e.g., in Ficus oleifolia, and in the neotropical Perebea humilis).

The abaxial sterile strip or groove, which is particularly clear in elongate inflorescences, indicates adaxial orientation of the flowers in essentially racemosely constructed inflorescences. Pistillate flowers tend to occur in the centre of bisexual (essentially cymosely constructed) inflorescences and staminate flowers in the periphery (Berg 1977).

Some flowers and inflorescences show clear adaptation to the mode of pollination. In other cases the primary modifications seem to be for the protection of developing stamens and ovaries against predation by insect larvae (cf. Berg 1990). This protection can often be related to insects breeding in staminate inflorescences before later pollination.

The inflorescences are mostly axillary. They may occur below the leaves on previous season’s growth, often so in deciduous species. Ramiflorous inflorescences are born on short-shoots which are often already present in the leaf axils and often continue to bear inflorescences below the leaves, even down to the smaller branches. Cauliflorous inflorescences on short spur-like branchlets or on branches with long internodes are found in Artocarpus and in Ficus. In the latter genus such branches produced at the base of the trunk may become stolon-like (flagelliflory or geocarpy). Cauliflory is also found in the African genus Treculia and the neotropical genus Clarisia.
Inflorescences typically in pairs, unisexual or bisexual, racemose, spicate, globose-capitate, capitate with a discoid to cup-shaped receptacle (and then with or without involucre), or with an urceolate receptacle, multi- to uniflorous, bracteate. — Moraceous flowers are small, monochlamydeous, and usually unisexual. Flowers with stamens and non-functional pistils are found in Ficus subg. Sycidium. The flower is often 4-merous, but the number of tepals may be more or less than 4, or the perianth may be absent (e.g. in Brosimum (male, America), in Hullettia (male and female?), Treculia (female, Africa), and Trilepisium (male, Africa), or strongly reduced as in several species of Ficus subg. Sycomorus sect. Sycocarpus (female, Asia).

The number of stamens is rarely more than 4 (e.g. in subg. Ficus), but often less; Artocarpus and many species of Ficus have only one. The tepals are either imbricate or valvate (as in Trophis), or narrow and not touching or overlapping each other (e.g. in Ficus subg. Synoecia). They may be free, or connate, forming tubular perianths, which can be fused with the ovary (e.g. in Trophis). Perianths of pistillate flowers either are or become ± fleshy; Fatoua and Ficus are notable exceptions.

The ovaries contain a single apically or subapically attached, anatropous to campylotropous ovule. There are two stigmata, or one by reduction. The stigmata vary in shape from filiform, to tongue-shaped, to truncate. Heterostyly is found in Ficus.

The stamens are straight in the bud in most genera, but in the majority of the species of the tribe Moreae inflexed and at anthesis bend outwards elastically. The anthers have two thecae but, due to fusion, seemingly only one in Ficus subsect. Malvanthera.

Pistillodes are found in flowers with the urticaceous type of stamens where they may play a role in keeping the anther in position. They are, moreover, characteristic for Ficus subg. Sycidium, in which they may be as large as pistils. Pistillodes are often found in flowers of Ficus, e.g. in subg. Pharmacosycea and only occasionally in other groups of Moraceae.
Flowers unisexual, free or connate (or also adnate to the receptacle). stamens 1-4(-6), straight or inflexed before anthesis; ovule 1, (sub)apically attached, anatropous to campylotropous. — The distribution of the family suggests that the dehiscent drupe might be the basic fruit type of Moraceae (Berg 1977). The exocarp is whitish, stipitate at the base, and laterally thickened. The turgid halves clasp the endocarp body, kept in its position by the vascular bundle running on top of it. When this bundle breaks the endocarp body is ejected from the fruit if it is small, or is squeezed out if it is large. It can also be ejected as in Streblus elongates and S. macrophylla, in the latter up to 10 m from the tree (pers. comm. Dr. D.J. Middleton). Small endocarp bodies, as can be found in Fatoua (and Dorstenia), are white and tuberculate; large ones are black and smooth (Antiaropsis, Streblus). Dehiscent drupes are found in Antiaropsis, Bleekrodea, Broussonetia, Dorstenia (except for the only epiphytic species, see Berg & Hijman 1999), some species of Ficus, Fatoua, Scyphosyce Baill. (Africa), Sparattosyce Bureau (New Caledonia), and Utsetela Pellegr. (Africa). Some of the features of the pericarp of the dehiscent drupe, such as the basally narrowed and/or laterally thicker pericarp, can also be found in indehiscent types.

The ovary is often fused with the perianth and the features described above are obliterated. Fruits with dry pericarps are found in the majority of the Ficus species. The stipitate base often found in the achene of Ficus might be homologous to the stipitate base of the dehiscent fruit.

The seed coat, protected by the more or less hard endocarp, is thin, but often conspicuously vascularised. Small seeds contain endosperm, as in Ficus, but in large seeds it is (almost) lacking. The embryos of small seeds are simple, those of large seeds more elaborate with thick, folded and/or unequally large cotyledons and short or long radicles.
Fruit an achene or drupaceous (dehiscent or indehiscent), free or adnate to the perianth, often forming a drupaceous whole with the fruiting perianth or also with the (fleshy) receptacle. Seed large without endosperm or small with endosperm;

Distribution

Africa: present America: present Asia: present Asia-Tropical:, Borneopresent; Jawa (Jawapresent); New Guineapresent; Sumatera (Sumaterapresent) Asian- Australasian region: present Australasia: present Madagascar: present Malesian region: present New Zealand and New Caledonia: present Pacific: present from West Africa to the Tonga Islands: present
The family comprises 37 genera and c. 1050 species represented by 14 native genera with in total 422 indigenous species; 8 species are introduced in Malesia of which 3 belong to Ficus.

The family is essentially tropical. Two genera, Broussonetia and Morus, are associated with and morphologically adapted to northern warm-temperate conditions. The genera Ficus and Maclura extend with a few members into warm-temperate conditions in the northern hemisphere and with two species of Streblus into those of the southern hemisphere.

Most species of Moraceae (c. 600 spp.) occur in the forest complex of tropical Asia and Australasia, where the large number of Ficus and, to a lesser extent, of Artocarpus swell the total. Africa with c. 185 species and America with c. 270 species are specifically poorer, but are richer in endemic genera: 7 out of 17 and 14 out of 19, respectively.

The third largest genus of the Moraceae, Artocarpus, is largely Malesian. About half of the species of the largest genus of the family, Ficus, occurs in the Malesian region.

The Asian element links closely to the Australasian, which can be regarded as an extension of the Asian. It is distinguished by the monotypic genus Sparattosyce, endemic to New Caledonia, and by some endemic or subendemic subdivisions of Ficus and Streblus.

Only six of the native genera, apart from Sparattosyce, are confined to the Asian- Australasian region. The other genera also occur in Madagascar, the African continent, and/or the Neotropics.

There are several links between the Asian moraceous flora and the African one. One pattern is formed by a number of subdivisions of Ficus (subsect. Pedunculatae of subg. Pharmacosycea, subsect. Urostigma of subg. Urostigma, sect. Sycidium of subg. Sycidium, and subsect. Ficus of subg. Ficus) which are centred in western Asian dry or seasonal forest but with linking elements in African savannah woodland. Another pattern is that between the Madagascan region, the eastern Asian mainland, and Malesia exhibited by the genera Bleekrodea, Broussonetia, Fatoua, Streblus, Trophis, and Ficus subg. Sycomorus sect. Sycomorus. Other links are formed by the close relationship between the African genus Treculia and the Asian genus Artocarpus and the African genus Milicia and Asian Moreae, and by Antiaris toxicaria ranging from West Africa to the Tonga Islands. Two other species occur in Asia and Africa: Ficus exasperata and F. palmata.

The links with America are much weaker and are realized by Ficus (subg. Pharmacosycea) and Trophis, which connect the north western part of the Neotropics to the eastern part of the Asian-Australasian region. The Neotropics and Africa are linked by the tribes Dorsteniae and Castilleae.

Ficus, Maclura, and Trophis are pantropical genera. The essentially northern warm-temperate genus Morus extends into the montane tropics of Asia and America and lowland Africa. Dorstenia, speciose in America and Africa, is represented in Asia by only one Indian species.

Within Malesia the greatest numbers of Moraceae coincide with those of the Dipterocarpaceae, except for the development of Ficus in New Guinea. Most genera (and subgenera of Ficus) have one or more widespread species. A Sino-Himalayan element infiltrates the Malay Peninsula and some of the species reach Sumatra, Java, and Borneo. An Australian element is evident in Ficus subsect. Malvanthera. It penetrates only the adjacent part of the Malesian region and the Pacific. The Australasian Streblus sect. Parastreblus extends to New Zealand and New Caledonia. The genera endemic to Malesia are Antiaropsis, restricted to New Guinea, and the more widespread Prainea. Parartocarpus slightly exceeds Malesia.

Two neotropical genera, Castilla and Dorstenia, have been introduced, each with one species. A few other species introduced in the Malesian region belong to Broussonetia, Ficus, and Morus and are of Asian origin.

Ecology

The majority of the species are elements of lowland, mostly evergreen forest or less frequently, seasonal forest. Trees of coastal, riparian, and secondary growth occur mainly in Ficus, but Streblus asper is characteristic of the more seasonal forest of western Malesia.

Some species of Ficus (F. deltoidea and F. oleifolia) often occur on poor sandy soil in kerangas forest. Other species of Ficus are associated with calcareous substrates (F. anastomosans, F. calcarata, F. calcicola, F. subcaudata (?), and the form F. tinctoria described as F. swinhoei). A form (ecotype) of F. ulmifolia is associated with extreme ultra basic soil (Philippines).

Hemi-epiphytic species can be hemi-epilithic on exposed rocky surfaces.

Some species of Ficus are more or less clearly rheophytic (see FM17-2: 27), e.g., F. ischnopoda, F. macrostyla, and F. squamosa, the latter two with fruitlets with long persistent styles and retrorse stiff hairs on the style and the margin of the fruit body morphologically adapted to the unusual substrate.

In general, the upper altitudinal limit of lowland species is 1500 m. Montane species, above 2000 m, are found in Ficus and Streblus. Morus macroura can also be regarded as montane. The hardier Moraceae of Sino-Himalaya do not reach Malesia.

Pollination

— Wind-pollination occurs in Moraceae with urticaceous type stamens and may occur in some other species, such as those with long pendulous staminate inflorescences. Anemophily is also reported for patent (scentless) staminate inflorescences of some species of Artocarpus (e.g., A. elasticus and A. rigidus (Corner 1940, 1988; Jarrett 1959) that give off clouds of pollen. The ballistic release of pollen from urticaceous type stamens allows plants to inhabit the forest undergrowth where they can make use of weak air currents, as above streaming water, for the transport of pollen. This type of pollination is described and discussed for Streblus pendulinus and Trophis scandens by Williams & Adam (1993).

Geitonogamy might occur in species with bisexual inflorescences, such as Dorstenia.

Insect-pollination is probably the predominant mode of pollination in the family. The unique mode in Ficus is well-documented (see FM 17-2: 51-54). Another is described for Artocarpus (Van der Pijl 1953), in which species with staminate inflorescences (e.g., A. dadah and A. integer; Corner 1940) emit a sweet scent of honey and burnt sugar, to attract small flies and beetles which subsequently breed in the inflorescence. Pollination based on insects breeding in staminate inflorescences is common in many tropical plant groups with reduced and unisexual flowers (such as palms), and is probably widespread in those Moraceae with dense unisexual inflorescences as well. In Castilla elastica and Antiaropsis decipiens this role is played by thrips (Sakai 2001; Zerega et al. 2004), which could be the case in other genera where the staminate inflorescences are (±) closed before anthesis. In some species of Artocarpus with scentless staminate inflorescences, clouds of pollen are given off, e.g. in A. elasticus and A. rigidus (Corner 1938, 1940; Jarrett 1959).

An unusual mode of pollination is described for A. integer in Sarawak (Sakai et al. 2000): staminate inflorescences infected and covered by fungus are visited by a species of gall midge that feeds on the mycelium and oviposits on the inflorescence. The midges transport pollen to pistillate inflorescences. It is not clear whether this is a local phenomenon or widespread in the species.

Dispersal

— The seeds are usually animal dispersed, although this may not be the case if the fruits are dehiscent drupes. Here, small endocarp bodies (kernels) are ejected (Dorstenia and Fatoua) or, if large, are squeezed out (Antiaropsis and Streblus p.p.), and drop on the forest floor or into water (by which they are carried on) or are ejected (see p. 8). The endocarp bodies, which are animal dispersed, remain inside the infructescence (Broussonetia and Ficus). Alternatively they may be large, black, and embedded in structures with contrasting colours (e.g. white of the exocarp and red of tepals as in Antiaropsis), in which case they may be dispersed by birds. In the majority of the Moraceae the fruits (and seeds) are enclosed in fleshy structures consisting of connate perianths and interfloral bracts (Artocarpus and related genera), or also adnate to the receptacle (Hullettia), or enclosed in fleshy receptacles (Ficus). The infructescence may more simply be just aggregations of flowers in fruit of which the fleshy perianths are the attractive parts (Morus). More or less solitary fleshy (dehiscent or indehiscent) fruits occur in Streblus. The colour of individual fruits or of infructescences varies from red to orange. Large infructescences are often greenish to yellowish. The infructescences of Ficus can be blackish, or sometimes brown or purplish and then taken by fruit bats.

Moraceous fruits and infructescences are taken by birds, monkeys, squirrels, and other arboreal animals by day. By night the same fruits, or those of allied species, are taken by nocturnal animals, such as fruit bats and civet cats. Cauliflorous species of Artocarpus and Ficus may also be dispersed by bats as well as by ground mammals, ranging from elephants and rhinoceros to pigs and mouse deer.

Water may play a role in distribution of diaspores ejected or fallen into streams. The fruitlets of some rheophytic Ficus species are adapted to attachment to the substrate.

Taxonomy

In the classification by Engler (1888) the family Moraceae included the subfamily Conocephaloideae, with six genera, including the Asian genus Conocephalus Blume. In a revisional study by Chew (1963) the genus was united with the urticaceous genus Poikilospermum Zipp. ex Miq. This led to proposals to transfer some or all the genera of the Conocephaloideae (Chew 1963 and Corner 1962, respectively) to the Urticaceae. For these six genera the family Cecropiaceae Berg (1978) was established.

The order Urticales constitute a clear-cut and coherent group. Moraceae differs from the Urticaceae in the presence of milky sap (or latex), the apical attachment of the ovule, the common presence of two stigmas, the absence of elongate cystoliths, and the predominantly woody habit in Urticaceae. The stamens are always inflexed in the bud and bend outwards suddenly, throwing the pollen into the air. Urticaceous stamens also occur in most species of the tribe Moreae. Inflexed stamens resembling those of Urticaceae occur in some genera of the Ulmaceae. The family Cecropiaceae is entirely woody and differs from the Moraceae in some features shared with Urticaceae, such as the basally attached ovule and single stigma, by strict dioecism, the absence of milky sap, the presence of adventitious roots (in the woody genera of Moraceae only present in Ficus), and the always spirally arranged leaves. Ulmaceae share with Moraceae the apically attached ovule, but lack milky sap. In contrast to the other families, the flowers are morphologically and often also functionally bisexual. Judd et al. (1994) suggested including Cannabidaceae, Cecropiaceae, Moraceae, in the Urticaceae, leaving the Celtidaceae and Ulmaceae as two much smaller urticalean families.

The Urticales show rather clear morphological affinities to the Malvales and somewhat remotely to the Euphorbiales, but they do not show links to the other families traditionally ranked among the Hamamelidae. Molecular studies place Urticales among the Rosales (Sytsma et al. 2002), but this contradicts patterns of morphological differentiation, ecology, and phytogeography, which suggest a different evolutionary history of the group (but see chapter ‘Phytochemistry and Chemotaxonomy’, p. 11).

For the reduced family Moraceae Corner (1962) proposed seven tribes, which with some adjustments was reduced to five tribes (Berg 1988, 2001): Moreae (8 genera, 70-75 spp., centred in Asia and characterized by the urticaceous type of stamen, see p. 7); Castilleae (8 genera, 50-60 spp., centred in the Neotropics and characterized by trees with the architectural model of Cook (see p. 138); Dorstenieae (8 genera, 125-130 spp., amphi-atlantic and characterized by circular bisexual inflorescences); and Ficeae (1 genus, 720-750 spp., characterized by essentially bisexual, urceolate inflorescences). More recently the tribe Artocarpeae was redefined and two additional tribes established (Berg 2005): Artocarpeae (4 genera, c. 55 spp, centred in Asia and characterized by many-seeded infructescences, mostly formed by connate flowers, but with free fruits); Antiaropsideae (2 genera, 3 or 4 spp.), in New Caledonia and New Guinea and characterized by involucrate inflorescences and dehiscent drupes); and Soroceae (5 genera, 23 spp., neotropical and characterized by simply constructed, Moreae-like inflorescences but with stamens straight in the bud).

In the classification of Moraceae proposed by Corner (1962) the number of genera previously recognised for the Asian-Australasian region was reduced considerably. This study and others by Berg (e.g., 1986, 1988), in which some of the Asian-Australasian representatives of the family were involved, resulted in the recognition of 37 genera worldwide. These appear to be largely natural ones and show interesting patterns in geographical distribution and taxonomic relationships.

A molecular phylogenetic study by Datwyler & Weiblen (2004) largely supports distinctness and homogeneity of Castilleae, Dorstenieae, and Ficeae, but not so of Artocarpeae (sensu stricto), Moreae, and Soroceae, not even at the generic level. Moreover, the study indicates affinities of Antiaropsideae to Castilleae and Ficeae. Support for this phylogeny from morphology and phytogeography is still wanting. The phylogeny of the more recent molecular study by Zerega et al. (2005) is rather similar and pays attention to biogeography and divergence times, but hardly to patterns in morphological differentiation.
G. Berg, C.C. - Flora Neotropica Monograph 83 in Moreae, Artocarpeae, Dorstenia (Moraceae). With introductions to the family and Ficus and with additions and corrections to Flora Neotropica Monograph 7. 2001, H. Berg, C.C., Cecropiaceae a new family of the Urticales in Taxon 27. 1978, I. Berg, C.C., Flora Malesiana precursor for the treatment of Moraceae 8: other genera than Ficus. in Blumea 50. 2005, J. Berg, C.C., The delimitation and subdivision of the genus Maclura (Moraceae) in Proc. Kon. Ned. Akad. Wetensch. 89. 1986, K. Berg, C.C., The genera Trophis and Streblus (Moraceae) remodelled in Proc. Kon. Ned. Akad. Wetensch. 91. 1988, L. Chew, W.-L., Flora malasianae precursores XXXIV A revision of the genus Poikilospermum (Urticaceae). in Gard. Bull. Singapore 20. 1963, M. Corner, E.J.H., The classification of Moraceae in Gard. Bull. Singapore 19. 1962, N. Datwyler, S.L. & G.D. Weiblen, On the origin of the fig: phylogenetic relationships of Moraceae from ndhF sequences in Amer. J. Bot. 91. 2004, O. Engler, G.H.A - Moraceae in Natürliche Pflanzenfamilien. 1888, P. Judd, W.S., R.W. Sanders & M.J. Donoghue, Angiosperm family pairs: preliminary phylogenetic analyses in Harvard Pap. Bot. 5. 1994, Q. Sytsma, K.J., J. Morawetz, J.C. Pires, M. Nepokroeff, E. Contoi, M. Zjhra, J.C. Hall & M.W. Chase, Urticalean rosids: circumscription, rosid ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences in Amer. J. Bot. 89. 2002, R. Zerega, N.J.C., W.L. Clement, S.L. Datwyler & G.D. Weiblen, Biogeography and divergence times in the mulberry family (Moraceae) in Molec. Phylogenet. Evol. 37. 2005

Cytology

There are two series, one with the haploid number of 13 and the other with 14 (Fedorov 1969; Oniguma & Tobe 1995). Brosimum, Broussonetia, Streblus p.p., and Ficus belong to the first series. Ficus are mostly diploids, but some (mainly African species) are or can be tetraploid, and F. elastica triploid (Ohri & Khoshoo 1987). The genera belonging to the second series are Antiaris, Artocarpus, Castilla, Clarisia, Maclura, Morus, Pseudolmedia, Scyphosyce (cf. Berg 1977), Streblus p.p., and Trophis. Some Arto-carpus species are tetraploid (or hexaploid in cultivars of A. altilis) and some Morus species polyploid, up to 2n = 308. Dorstenia shows much variation in its chromosome numbers: 2n = 24, 26, 28, 30, 32, 36, 38, 40, 42, 48, 52, and c. 64 (Berg & Hijman 1999). Deviating numbers are reported for Maclura tricuspidata, 2n = 50 (Morawetz & Samuel 1989) and Naucleopsis guianensis, 2n = 20 (Dmitrieva & Parfenov 1985). The haploid chromosome number of both Cecropiaceae and Ulmaceae is 14, that of Urticaceae varies from 7 to 14. The chromosome numbers of the Cannabidaceae deviates from the general pattern in the order.
S. Berg, C.C. & M.E.E. Hijman, The genus Dorstenia (Moraceae) in Ilicifolia 2. 1999, T. Berg, C.C., Revisions of African Moraceae (excluding Dorstenia, Ficus, Musanga and Myrianthus) in Bull. Jard. Bot. État 47. 1977, U. Chromosome numbers of flowering plants. 1969, V. Dmitrieva, S.A. & V.I. Parfenov, Kariologicheskaja kharakteristika nekotorykhvidov poleznykh rastenij flori Belorussii in Izvestii Akademii Nauk Belorusskoi SSR: Seriia Biologicheskikh Nauk 6. 1985, W. Morawetz, W. & M.R.A. Samuel - Karyological patterns in Hamamelidae in Evolution, systematics and fossil history of the Hamamelidae: ‘Introduction’ and ‘lower’ Hamamelidae. 1989, X. Ohri, D. & T.N. Khoshoo, Nuclear DNA contents in the genus Ficus (Moraceae) in Pl. Syst. Evol. 156. 1987, Y. Oniguma, K. & H. Tobe, Karyomorphology of some Moraceae and Cecropiaceae (Urticales) in J. Pl. Research 108. 1995

Uses

Streblus elongatus and several species of Artocarpus are, or have been, outstanding producers of timber. Lesser known are the timber trees found in the genera Antiaris, Parartocarpus, and Prainea (Boer & Sosef 1998). Ficus elastica and the American Castilla elastica have been used for rubber and remains of their plantations may be found throughout Malesia, but they have given place to Hevea because of their resinous milksap. Poisonous milksap occurs in the famous upas tree Antiaris (see Boer et al. 1999) and in the much less common Parartocarpus; that of both is used as (one of the components of) dart and arrow poison in South-east Asia. The sap of many species of Artocarpus, by contrast, being very sticky and innocuous, is used for birdlime and general adhesive; whereas that of A. lowii, is oily and greasy and is used as an ointment and for cooking. Fibrous and easily-stripped bark of several species of Artocarpus, such as A. elasticus, is made into coarse bark cloth and binding material, but this is disappearing even from the life of jungle folk; many anthropological exhibits in museums are held together by this material. The fibrous bark of Ficus is used for string, even for bow strings, but being readily cropped these plants have not been commercialised. Paper is manufactured from the inner bark of the paper mulberry Broussonetia papyrifera, as well as fibre clothing and string among primitive people (Berg 2003). Edible fruits are the mulberries (Morus), various kinds of figs (Ficus), several kinds of Artocarpus, and indeed Antiaris, the fruits of which seem always devoid of poison. Introduced mulberries have found little favour in Malesia. The cultivated fig (Ficus carica) is rarely seen because it succumbs readily to insect attacks. Sometimes the Indochinese Ficus auriculata is cultivated, but its fruit is inferior. Certain wild figs are, however, not despised. The best known fruit trees of the family in Malesia are the jackfruit (Artocarpus heterophyllus) and the chempedak or chemedak (A. integer). They are grown both for the edible pulp around the seeds and for the seeds themselves which are roasted. Some varieties of the jack produce infructescences, which may hold the world record for fruit size. Few fruits can equal in stench some varieties of the chempedak. The breadfruit tree (Artocarpus altilis), commonly seen in villages, is extremely abundant both wild and cultivated in New Guinea, where it is grown for the unripe fruit which is baked, roasted, or boiled more like a tuber than a fruit; the seedless form is most usually cultivated. Artocarpus odoratissimus is cultivated in Borneo and the Philippines for the sweet pulp round the seed. Artocarpus nitidus, with the uniformly succulent kind of infructescence, is sometimes found in gardens in West Malesia, where it is used also for jam and conserves. Other species of Artocarpus are mostly wild trees which have been spared from felling for sake of their fruit. Leaves and young figs of several Ficus species are eaten either cooked or raw as vegetables.

Yellow dye is extracted from wood of Maclura cochinchinensis (Heyne 1927).
Z. Berg, C.C. - Broussonetia papyrifera (L.) L’Hér. ex Vent. in Plant Resources of South-East Asia , Fibre plants. 2003, AA. Boer, E. & M.S.M. Sosef - Antiaris, Parartocarpus, and Prainea in Timber trees: Lesser known timbers. Plant Resources of South-East Asia. 1998, AB. Boer, E., M. Brink & M.S.M. Sosef - Antiaris toxicaria Lesch. in Medicinal and poisonous plants 1. Plant Resources of South-East Asia. 1999, AC. Heyne, K., De nuttige planten van Nederlandsch-Indië in - undefined journal - Ruygrok & Co., Batavia. 1927

Phytochemo

Relatively few members of the family have been examined thoroughly by phytochemists. So the chemical characterisation must be regarded as a very incomplete and provisional one. The following features seem to be typical to some extent of Moraceae:
  1. Most members of the family deposit large amounts of silicic acid and calcium carbonate in the walls of leaf cells, especially hairs and unspecialised epidermal cells. Cystoliths and crystals of calcium oxalate are ubiquitous.
  2. Mucilage cells and ducts occur frequently. Thorough chemical characterisation of different kind of mucilage of Moraceae are still lacking.
  3. Latex-cells and non-articulated latex tubes are widespread. Some species produce rubber-rich latex. In other species, ‘resins’ (triterpenic alcohols and their acetates and cinnamates) or ‘waxes’ (ethers of cerotinic acid, esters of triterpenic alcohols with several fatty acids) predominate. Still other species produce protein-rich latex; ficin is a papain-like enzyme from the latex of several species of Ficus, ficin containing latex is used locally as anthelmintic (in the Neotropics).
    A few genera are known to contain cardio-toxic compounds; these have been isolated from, or demonstrated to be present in, latex of Antiaris, Castilla, Naucleopsis (= Ogcodeia), barks of Streblus, and seeds of Antiaris, Antiaropsis, Castilla, and Naucleopsis. The latex of Antiaris toxicaria is used as arrow poison in eastern Asia, and that of some species of Naucleopsis for the same purpose in the Neotropics (Bisset & Hylands 1977). Still other species seem to produce latexes which are very rich in phenolic compounds; Vreede (1949) observed flavonoids (not definitely defined) in yellow latexes of several species of Ficus. Chlorogenic acid was isolated from the latex of Castilla elastica.
  4. Polyphenolic compounds are common in Moraceae. Derivates of p-coumaric acid, caffeic acid, kaempferol, and quercetin occur frequently in leaves. Myricetin is less common. Leucoanthocyanins are absent of the leaves of many species, but often present in species of Ficus. The so-called tannin-idioblasts of the anatomical literature are present in the mesophyll of many species. They probably contain catechins and leucoanthocyanins, but no true tannins, which have yet to be demonstrated in Moraceae for certain.
  5. The compounds of most interest from the systematic point of view are the highly characteristic phenolic compounds which seem to be rather common in the roots, stems, fruits, and sometimes the leaves. These phenolics may roughly be classified in four groups: C6-C3-compounds (coumarins), represented by furanocoumarins and dimethylpyranocoumarins); C6-C1-C6-compounds (benzophenones like maclurin and xanthones); C6-C2-C6-compounds (stilbenes like chlorophorin and hydroxyresveratrol); C6-C3-C6-compounds (highly characteristic flavonoids like cyanomaclurin, morin, artocarpetin, cycloartocarpin, and pomiferin). The resistance of the woods of some members of the family to the attack by fungi, insects, and termites, and the tinctorial properties of the woods of others, are largely due to such polyphenolic compounds.
  6. Saponins and alkaloids seem to be rather rare in Moraceae. Alkaloids are known from some species of Ficus. No taxonomic implications are apparent at present from these occurrences.


To sum up, the accumulation of minerals in leaves (SIO2, CaCO3), and the production of a whole array of unusual phenolic compounds represent the most striking, currently known, phytochemical features of Moraceae.

With regard to the different mineralizations (including the presence of cystoliths) the family resembles Cannabidacea and Urticaceae.

Some outstanding features of phenolic compounds of Moraceae indicate similarities in secondary metabolism with members of the rosalean alliance, especially with Leguminosae.

These features are: production of furanocoumarins, isoflavones (osajin, pomiferin), stilbenes, flavonoids with a resorcin-type hydroxylation pattern of the B-ring (cyanomaclurin, artocarpetin, morin, etc.), and the frequent attachment of isoprenoid substituents to aromatic rings (coumarins, stilbenes, and flavonoids).

Isoprenylation also marks moraceous xanthones (alvaxanthone, macluraxanthone), in which respect the family resembles Guttiferae. Knowledge of the distribution of all these types of phenolics is still rather scanty. It would be cautious therefore not to put too much weight on the metabolic similarities mentioned. A more comprehensive review of chemical characters may be found in Hegnauer (1969).

Volatile compounds produced by inflorescences of Ficus play an important role in attraction of pollinators. Compounds extracted from receptive syconia of F. carica include benzyl alcohol, linanool, linanool oxides (furanoid), several aromatic compounds (cinnamic aldehyde, cinnamic alcohol) and indole (Gibernau et al. 1997).
AD. Bisset, N.G. & P.J. Hylands, Cardiotonic glycosides from latex of Naucleopsis mello-barretoi, a dart poison plant from north-west Brazil in Econ. Bot. 31. 1977, AE. Gibernau, M., H.R. Buser, J.E. Frey & M. Hossaert-McKey, Volatile compounds from extracts of figs of Ficus carica in Phytochemistry 46. 1997, AF. Hegnauer, R., Chemotaxonomie der Pflanzen in - undefined journal - 5. 1969, AG. Vreede, M.C., Topography of the lactiferous system in the genus Ficus in Ann. Bot. Gard. Buitenzorg 51. 19491. R. Hegnauer

Citation

Benth. & Hook.f. - in Gen. Pl. 1880: (Urticaceae)
Link - in Proc. Kon. Ned. Akad. Wetensch. 1988
Cronquist, An integrated system of classification of flowering plants. 1981
Corner - in Gard. Bull. Singapore. 1962
Baill. - in Hist. Pl. 1875: (Ulmacées)
C.C. Berg - in Taxon. 1978
Engl. - in Engl. & Prantl, Nat. Pflanzenfam. 3. 1888