Family: Moraceae

Broussonetia papyrifera
Broussonetia papyrifera   (L.) L¥HÈr ex Vent.  -  Paper Mulberry
Photo © by Steven Clemants
Taken in Prospect Park, Brooklyn, NY, 1996.

By Katherine Gould, Angela Steward, & Steven D. Glenn

Not peer reviewed

Last Modified 03/04/2013

Key to the genera of Moraceae

1. Leaves entire, pinnately veined; stems usually thorny; fruit a large globose aggregate...Maclura
1. Leaves toothed or lobed, palmately veined; stems not thorny; fruit various...2

2. Terminal vegetative bud surrounded by a pair of stipules...Ficus
2. Terminal vegetative bud not surrounded by a pair of stipules...3

3. Leaves densely hairy on blade and veins beneath; fruit globose, inedible...Broussonetia
3. Leaves glabrous, scabrous, or sparsely hairy on veins beneath; fruit short-cylindric, edible, and resembling a blackberry...Morus

List of Moraceae Genera

References to Moraceae

  • Angelo, R.; Boufford, D. E. 2010. Atlas of the flora of New England: Magnoliidae and Hamamelidae. Rhodora 112: 244-326.
  • Bechtel, A. R. 1921. The floral anatomy of the Urticales. Amer. J. Bot. 8: 386-410.
  • Beck, N. G.; Lord, E. M. 1988. Breeding system in Ficus carica, the common fig: 1. Floral diversity. Amer. J. Bot. 75(12): 1904-12.
  • Beck, N. G.; Lord, E. M. 1988. Breeding system in Ficus carica, the common fig: 2. Pollination events. Amer. J. Bot. 75(12): 1913-22.
  • Berg, C. C. 1989. Classification and distribution of Ficus. Experientia (Basel) 45(7): 605-11.
  • Berg, C. C. 1986. The delimitation and subdivision of the genus Maclura (Moraceae). Proc. Kon. Nederl. Akad. Wetensch., C, 89(3): 241-7.
  • Berg, C. C. 1990. Reproduction and evolution in Ficus (Moraceae): traits connected with the adequate rearing of pollinators. Mem. New York Bot. Gard. 55: 169-85.
  • Braun, J.; Brooks, G. R. 1987. Box turtles (iTerrapene carolina) as potential agents for seed dispersal. Amer. Midl. Naturalist 117: 312-8.
  • Bronstein, J. L.; McKey, D. 1989. The comparative biology of figs. The fig/pollinator mutualism: a model system for comparative biology. Experientia (Basel) 45(7): 601-4.
  • Bryant, F. C. 1996. Diets of female white-tailed deer in the cross-timbers region. Prairie Naturalist 28: 125-140.
  • Burgess, K. S.; Husband, B. C. 2004. Maternal and paternal contributions to the fitness of hybrids between red and white mulberry (Morus, Moraceae). Amer. J. Bot. 91: 1802-1808.
  • Burton, J. 1973. Osage-orange: an American wood. USDA Forest Service.
  • Burton, P. J.; Bazzaz, F. A. 1991. Tree seedling emergence on interactive temperature and moisture gradients and in patches of old-field vegetation. Amer. J. Bot. 78(1): 131-49.
  • Carr, M. E. 1985. Plant species evaluated for new crop potential. Econ. Bot. 39(3): 336-45.
  • Clement, W. L.; Weiblen, G. D. 2009. Morphological evolution in the mulberry family (Moraceae). Syst. Bot. 34: 530-552.
  • Collingwood, G. H. 1939. Osage-orange. Amer. Forests 45: 508-10.
  • Condit, I. J. 1932. The structure and development of flowers in Ficus carica L. Hilgardia 6: 443-81.
  • Connolly, B. A. 2008. Six new vascular plant taxa for Connecticut. Rhodora 110: 354-358.
  • Corner, E. J. H. 1962. The classification of Moraceae. Gard. Bull. Straits Settlem. 19: 187-252.
  • Cowart, N. M.; Graham, J. H. 1999. Within- and among-individual variation in fluctuating asymmetry of leaves in the fig (Ficus carica L.). Int. J. Plant Sci. 160: 116-121.
  • Crane, P. R.; Blackmore, S. (eds.) (1989): 1989. Evolution, systematics and fossil history of the Hamamelidae. 2 Vols. Oxford University Press, New York.
  • Cross, G. L. 1937. The origin and development of the foliage leaves and stipules of Morus alba. Bull. Torrey Bot. Club 64: 145-63.
  • Cross, G. L. 1936. The structure of the growing point and the development of the bud scales of Morus alba. Bull. Torrey Bot. Club 63: 451-65.
  • Datwyler, S. L.; Weiblen, G. D. 2004. On the origin of the fig: phylogenetic relationships of Moraceae from NDHF sequences. Amer. J. Bot. 91: 767-777.
  • Debor, H. W.; Grosse, B. 1977. Bibliography of the international literature on the fig (Ficus sp.). (Ref. in Bibliogr. Agric., 41(12):120149. 1977.)
  • Duke, J. A. 1983. The handbook of engery crops.
  • Duke, J. A.; Ayensu, E. S. 1985. Medicinal plants of China. 2 Vols. Reference Publications, Algonac, MI. , 705 pages.
  • Flora of North America Editorial Committee. 1997. Flora of North America, Volume 3. Magnoliophyta: Magnoliidae and Hamamelidae. Oxford University Press, New York. , 590 pages.
  • Galil, J.; Neeman, G. 1977. Pollen transfer and pollination in the common fig (Ficus carica L.). New Phyt. 79(1): 163-71.
  • Galla, S. J. 2009. Morus murrayana (Moraceae): a new mulberry from eastern North America. Phytologia 91: 105-116.
  • Gray, E. 1990. Evidence of phenotypic plasticity in Mulberry (Morus L.). Castanea 55: 272-81.
  • Gray, E.; Call, N. M. 1994. Effects of induced plant injury on leaf lobation in red mulberry (Morus rubra L.). Castanea 59: 167-75.
  • Gray, E.; Gray, R. E. 1987. Leaf lobation patterns in mulberry. Castanea 52: 216-24.
  • Greive, M. 2000. A modern herbal.
  • Gupta, B. K.; Bhakuni, B. S. 1991. Inflorescence behavior of mulberry. Indian J. Forest. 14: 251-2.
  • Hans, A. S. 1972. Cytomorphology of arborescent Moraceae. J. Arnold Arbor. 53: 216-25.
  • Hardin, J. W. 1981. Atlas of foliar surface features in woody plants, II. Broussonetia, Morus, and Maclura of North America. Bull. Torrey Bot. Club 108: 338-46.
  • Herre, E. A. et al. 1996. Molecular phylogenies of figs and their pollinator wasps. J. Biogeogr. 23: 521-30.
  • Jennings, O. E. 1920. The paper mulberry (Broussonetia), an "artillery plant". Torreya 20: 52-3.
  • Khadari, B. et al. 1995. When figs wait for pollinators: the length of fig receptivity. Amer. J. Bot. 82(8): 992-9.
  • Kim, M.; Zavada, M. S. 1993. Pollen morphology of Broussonetia (Moraceae). Grana Palynol. 32: 327-9.
  • Kjellberg, F. et al. 1987. The stability of the symbiosis between dioecious figs and their pollinators: a study of Ficus carica L. and Blastophaga psenes L. Evolution 41: 693-704.
  • Kjellberg, F.; Maurice, S. 1989. Seasonality in the reprodutive phenology of Ficus: its evolution and consequences. Experientia (Basel) 45(7): 653-60.
  • Krefting, L. W.; Roe, E. I. 1949. The role of some birds and mammals in seed germination. Ecol. Monogr. 19: 269-286.
  • LeCoq, C. 1963. Contribution a l'etude cytotaxonomique de Moracees et de Urticacees. Rev. Gen. Bot. 70: 385-426.
  • McVaugh, R. 1952. Suggested phylogeny of Prunus serotina and other wide ranging phylads in North America. Brittonia 7: 317-346. (And other genera)
  • Mears, J. A. 1973. Chemical constituents and systematics of Amentiferae. Brittonia 25(4): 385-94.
  • Miller, N. G.; Wood, C. E. 2003. The Asian weed Fatoua villosa (Moraceae) in New York State and Massachusetts. Rhodora 105: 286-291.
  • Mitchell, R. S. (eds.) (1988): 1988. Platanaceae through Myricaceae of New York State. New York State Museum Bull. No. 464. The University of the State of New York, the State Education Department, Albany. , 98 pages.
  • Morton, J. 1987. Fruits of warm climates.
  • Nakai, T. 1927. Morus alba and its allies, in the herbaria of Linnaeus, Thunberg and others. J. Arnold Arbor. 8: 234-8.
  • Neeman, G.; Galil, J. 1978. Seed set in the 'male syconia' of the common fig Ficus carica L. (Caprificus). New Phyt. 81(2): 375-80.
  • Pair, J. C. 1992. Magnificent Maclura- past and present. Arnoldia (Jamaica Plain) 52: 14-9.
  • Penny, G. M.; Neal, J. C. 2003. Light, temperature, seed burial, and mulch effects on mulberry weed (Fatoua villosa) seed germination. Weed Technol. 17: 213-218.
  • Petersen, F. P.; Fairbrothers, D. E. 1985. A serotaxonomic appraisal of the "Amentiferae". Bull. Torrey Bot. Club 112: 43-52.
  • Philpott, J. 1953. A blade tissue study of leaves of forty-seven species of Ficus. Bot. Gaz. 115(1): 15-35.
  • Qui, Y. L. 1998. Phylogenetics of the Hamamelidae and their allies: parsimony analysis of nucleotide sequences of the plastid gene rbcL. Int. J. Plant Sci. 159: 891-905.
  • Ramirez, B. W. 1978. A new classification of Ficus. Annual Rep. Missouri Bot. Gard. 64(2): 296-310.
  • Rao, C. K.; Jarvis, C. E. 1986. Lectotypification, taxonomy and nomenclature of Morus rubra, M. tatatica and M. indica (Moraceae). Taxon 35: 705-8.
  • Rehder, A. A. 1945. Moraceae, Hippocastanaceae et Vitaceae, nomina conservanda. J. Arnold Arbor. 26: 277-9.
  • Robinson, B. L. 1907. The scientific name of the Osage orange. Rhodora 9: 91.
  • Sand, S. 1991. A tree history- the osage orange. Amer. Horticulturist 70: 37-39.
  • Sanford, S. N. F. 1933. Ficus carica in Massachusetts. Rhodora 35(409): 40.
  • Schnabel, A.; Laushman, R. H.; Hamrick, J. L. 1991. Comparative genetic structure of two co-occurring tree species, Maclura pomifera (Moraceae) and Gleditsia triacanthos (Leguminosae). Heredity 67: 357-64.
  • Schneider, S. 1998. Red mulberry: Morus rubra.
  • Seymour, F. C. 1952. Notes on Moraceae and Ulmaceae. Amer. Midl. Naturalist 48: 249-50.
  • Smith, J. L.; Perino, J. V. 1981. Osage orange (Maclura pomifera): history and economic uses. Econ. Bot. 35(1): 24-41.
  • Stafford, P. J. 1988. Evolution, systematics, and fossil history of the Hamamelidae.
  • Stapanian, M. A. 1982. A model for fruiting display: seed dispersal by birds for mulberry trees. Ecology 63(5): 1432-43.
  • Steen, D. A. et al. 1995. The big trees of Michigan: 12. Morus rubra L. Michigan Bot. 34: 147-9.
  • Steffey, J. 1980. The mulberry family. Amer. Horticulturist 59(3): 12-3.
  • Stern, W. L. 1973. Development of the amentiferous concept. Brittonia 25(4): 316-33.
  • Stone, D. E. 1973. Patterns in the evolution of amentiferous fruits. Brittonia 25(4): 371-84.
  • Sullivan, J. 1993. Morus rubra. ()
  • Thorn1. 1973. The "Amentiferae" or Hamamelidae as an artificial group: a summary treatment. Brittonia 25(4): 395-405.
  • Tiffney, B. H. 1986. Fruit and seed dispersal and the evolution of the Hamamelidae. Ann. Missouri Bot. Gard. 73: 394-416.
  • Tippo, O. 1938. Comparative anatomy of the Moraceae and their presumed allies. Bot. Gaz. 100: 1-99.
  • Valdeyron, G.; Lloyd, D. G. 1979. Sex differences and flowering phenology in the common fig, Ficus carica L. Evolution 33(2): 673-85.
  • Venkataraman, K. 1972. Wood phenolics in the chemotaxonomy of the Moraceae. Phytochemistry 11: 1571-86.
  • Verkerke, W. 1989. Structure and function of the fig. Experientia (Basel) 45(7): 612-22.
  • Vincent, M. A. 2004. Spread of Fatoua villosa (mulberry weed; Moraceae) in North America. J. Kentucky Acad. Sci. 65: 67-74.
  • Ware, A. B.; Kaye, P. T.; Compton, S. G.; Van Noort, S. 1993. Fig volatiles: their role in attracting pollinators and maintaining pollinator specificity. Pl. Syst. Evol. 186: 147-56.
  • Werner, P. A.; Harbeck, A. L. 1982. The pattern of tree seedling establishment relative to staghorn sumac cover in Michigan old fields. Amer. Midl. Naturalist 108: 124-32.
  • Wiebes, J. T. 1979. Co-evolution of figs and their insect pollinators. Ann. Rev. Ecol. Syst. 10: 1-12.
  • Wiegrefe, S. J.; Sytsma, K. J.; Guries, R. P. 1998. The Ulmaceae, one family or two? Evidence from chloroplast DNA restriction site mapping. Pl. Syst. Evol. 210: 249-270.
  • Yamazaki, T. 1982. The seed formation of Fatoua villosa Moraceae. Jap. J. Bot. 57: 358-365. (In Japanese)
  • Zavada, M. S.; Kim, M. 1996. Phylogenetic analysis of Ulmaceae. Pl. Syst. Evol. 200(1-2): 13-20. (also considers Moraceae & Cannabaceae)