Smilacaceae Vent.

Smilacaceae, E.J. Cowley. Flora of Tropical East Africa. 1989

Morphology General Habit

Robust, climbing or pendent shrubs, usually dioecious, glabrous, rarely pubescent; roots arising from compact rhizome

Morphology Stem

Stems and branches often aculeate

Morphology Leaves

Leaves alternate or opposite, petiolate, thin or coriaceous, curvinerved, with 3-7 main nerves Modified stipules forming 2 caducous tendrils arising from distinct, persistent leaf-sheath at base of petiole

Morphology Reproductive morphology Inflorescences

Inflorescence of few-many-flowered umbels; umbels solitary, or arranged in racemes or spikes

Morphology Reproductive morphology Flowers

Flowers regular, unisexual; perianth-segments 6, free or rarely united Female flowers with 1-6 filiform staminodes; ovary superior, sessile, 3-locular; locules with 1-2 ovules; stigmas 3, recurved Male flowers with 3 or 6(-15) stamens; filaments sometimes united into a column; anthers introrse; pistillode absent

sex Male

Male flowers with 3 or 6(-15) stamens; filaments sometimes united into a column; anthers introrse; pistillode absent

sex Female

Female flowers with 1-6 filiform staminodes; ovary superior, sessile, 3-locular; locules with 1-2 ovules; stigmas 3, recurved

Morphology Reproductive morphology Fruits

Fruit a 1-3-seeded, globose or broadly obovoid berry

Morphology Reproductive morphology Seeds

Seed globose or lenticular; endosperm hard

Botina-Papamija, J.R. (2009). Neotropical Smilacaceae.

Morphology

Description

Vines, usually climbing by paired tendrils, or rarely erect herbs, often with thick, tuber -like rhizomes. Stems rounded or sometimes quadrangular, armed with spines or unarmed , the surface smooth, scabrous , villose or setose . Base of branchlets provided with one or two scales on the adaxial side. Leaves simple , alternate , petiole usually geniculate, with sheath terminating in a pair of tendrils; blade broadly cordate to lanceolate , glabrous , smooth or ciliate , spiny or with unbranched hairs, with 3-9 subparallel primary  veins arising from or near the base (3 inner veins ), the lower order of venation reticulate , the margin entire . Inflorescences pedunculate, axillary , an umbela or panicle of umbels. Flowers unisexual (the plants dioecious ); perianth segments 6, distinct, narrow, equal or nearly so; stamens 6, free , the anthers basifixed, 2-celled, dehiscing longitudinally; ovary superior , syncarpous, 1- or 3-locular, with 1 or 2 ovules in each locule , the placentation axile ; stigmas 3, sessile , in female flowers 0-6 staminodes present. Fruit a fleshy and globose berry , indehiscent , smooth, orange, yellow, or black when mature; seeds 1-3, arillate, less than 10 mm in size.

General Description

Notes on delimitation

• The family Smilacaceae is placed by the APG II in the Liliales, together with four families which also occur in the Neotropics: Alstroemeriaceae, Corsiaceae, Liliaceae, and Melanthiaceae.

Number of genera

• The family is represented in the Neotropics by the genus Smilax  L. only, comprising ca. 100 species.

Status

• Smilax is native to the Neotropics.

General notes

• Smilacaceae has often been considered to be related to Dioscoreaceae, order Dioscoreales, but actually there is agreement among botanists that the family belongs in the Liliales. The family has also been included in the polyphyletic Liliaceae s.l. Since the publication of APG II (2003) it is accepted that Smilacaceae comprises two genera, Smilax and Heterosmilax Kunth, and ca. 300 species worldwide. However, some botanists have recently considered the family to comprise only Smilax with ca. 200 species.
• The family, and specially Smilax, is distributed in tropical, subtropical, and temperate areas of both hemispheres, with most species concentrated in the Neotropics and tropical Asia.
• In the Neotropics the rhizomes of most species of Smilax are used medicinally, but especially S. officinalis Kunth, S. longifolia Rich., S. siphilitica Humb. & Bonpl. ex Willd., S. aristolochiifolia Mill., S. moranensis M.Martens & Galeotti and S. domingensis Spreng. ex A. DC.  The stems of some species such as S. tomentosa Kunth are used in basket-making.  

Distribution

Distribution in the Neotropics

• The family Smilacaceae is distributed throughout the Neotropics, occupying several habitats at elevations from 0 to 3,200 meters.

Diagnostic

Distinguishing characters (always present)

• Leaves simple, alternate, with tendrils positioned at end of the petiolar sheaths.
• Leaf blade with 3-9 subparallel primary veins and reticulate secondary veins.
• Inflorescences axillary, an umbela or panicle of umbels.
• Flowers unisexual, dioecious.
• Perianth segments 6, distinct.
• Stamens 6, free.
• Fruit a fleshyberry with 1-3 seeds.

Other important characters

• Climbing vines.
• Petiole geniculate.

Key differences from similar families

Smilacaceae is similar to Dioscoreaceae in comprising vines with net-veined leaves. However, Smilacaceae has:

• Leaves with petiolar sheaths terminating in a pair of tendrils.
• Inflorescences an umbel or panicle of umbels.
• Fruit a berry with 1-3 seeds.

 Whereas family Dioscoreaceae has:

• Leaves without petiolar sheaths nor tendrils.
• Inflorescences mostly spicate, racemose, or paniculate.
• Fruit a 3-winged loculicidal capsule.

Useful tips for generic identification

Smilax is easily recognized by:

• Reticulate leaf venation.
• Paired petiolar tendrils.
• Dioecious flowers.
• Inflorescences an umbel or panicle of umbels.
• Fruit a berry.
• Climbing habit.

Literature

Important literature

Andreata, P.R.H. 1997. Revisão das espécies brasileiras do gênero Smilax Linnaeus (Smilacaceae). Pesq. 47: 1-243.

APG II. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Bot. J. Linn. Soc. 141: 399-436.

Botina-P., J.R. 2008. Revisión taxonómica y fitogeografía del género Smilax L.(Smilacaceae) en Colombia. Trabajo de Investigación. Universidad del Valle, Facultad de Ciencias Exactas y Naturales, Departamento de Biología, Programa Maestría en Ciencias-Biología, Santiago de Cali-Colombia.

Cameron, K.M. and Fu, C.X. 2005. A nuclear rDNA phylogeny of Smilax (Smilacaceae). Aliso 22: 598-605.

Chen, S.C., Qiu, Y.X., Wang, A.L., Cameron, K.M. and Fu, C.X. 2006a. A phylogenetic analysis of the Smilacaceae based on morphological data. Act. Phytotax. Sinica 44(2): 113-125.

Chen, S.C., Zhang, S.P., Ni, S.F., Fu, C.X. and Cameron, K.M. 2006b. The systematic value of pollen morphology in Smilacaceae. Pl. Syst. Evol. 259: 19-37.

Conran, J.G. 1998. Smilacaceae. Pp. 417-422. In: K. Kubitzki (ed.), The families and Genera of Vascular Plants, vol. 3.  Springer Verlag, Berlin.

Cronquist, A. 1981. An integrated system of classification of flowering plants. Columbia University Press, New York.

Dahlgren, R.M.T., Clifford H.T. and Yeo, P.E. 1985. The families of monocotyledons. Springer-Verlag, Berlin.

De Candolle, A.P.P. 1878. Smilax. Pp. 45-213. In: De Candolle, A.P.P. and C.P. De Candolle (eds.), Monographiae Phanerogamarum. Sumptibus G. Masson, Parisiis.

Ferrufino, L. and Gómez, J. 2004. Estudio morfológico de Smilax L. (Smilacaceae) en Costa Rica, con implicaciones sistemáticas. Lankesteriana 4(1): 5-36.

Gaskin, J. F. and Berry, P.E. 1988. New synonymy and useful taxonomic characters in Smilax (Smilacaceae) from the Venezuelan Guayana. Novon 8: 364-370.

Gaskin, J.K. and Berry, P.E. 2005. Smilacaceae. In: J.A. Steyermark, P.E. Berry, K. Yatskievich and B.K. Holst (eds.), Flora of the Venezuelan Guayana, vol. 9, Rutaceae -Zygophyllaceae. Missouri Botanical Garden, St. Louis, Missouri.

Grisebach, H.A. 1842. Smilaceae. Pp. 1-24. In: C.F.P. von Martius et al. (eds.), Flora Brasiliensis, vol. 3, n 1, Leipzig, München.

Guaglianone, E. R. and S. Gattuso. 1991. Estudios taxonómicos sobre el género Smilax (Smilacaceae) I. Bol. Soc. Argent. Bot. 27(1-2): 105-129.

Howard, R. A. 1979. Genus Smilax L. in the Lesser Antilles. Taxon 28: 55-58.

Huft, M. J. 1994. Smilacaceae. Pp. 20-25. In: G. Davidse, M. Sousa & A.O. Chater (eds.), Flora Mesoamericana, Vol. 6, Alismataceae a Cyperaceae. Universidad Autónoma de México, México D.F.; Missouri Botanical Garden, St. Louis; The Natural History Museum, London.

Judd, W. S., Campell, C.S., Kellogg, E.A., Stevens, P.F. and Donoghue, J.M.  2002. Plant systematics, a phylogenetic approach. 2 ed. Sinauer Associates, Inc., Sunderland, Massachusetts.

Killip, E.P. and Morton, C.V. 1936. A revision of the Mexican and Central American species of Smilax. Carnegie Inst. Wash. Publ. 461: 255-296.

Kunth, C.S. 1850. Smilax. Pp. 160-263 in Enumeratio Plantarum, Tomus V. Sumtibus J. G. Cottae, Stutgardiae et Tubingae.

Maas, P.J.M. and Westra, L.Y.T. 2004. Neotropical plant families. 3rd ed. A.R.G. Gantner Verlag K.G., Vaduz, Liechtenstein.

Morales, F. J. 2003. Smilacaceae. Pp. 833-838. In: B.E. Hammel, M.H. Grayum, C. Herrera and N. Zamora (eds.), Manual de Plantas de Costa Rica, vol. 3, Monocotiledoneas. Monogr. Syst. Bot. Missouri Bot. Gard. 93.

Morton, C.V. 1945. Smilacaceae. Pp. 6-11. In: R. E. Woodson and R.N. Schery (eds.), Flora of Panama. Ann. Missouri Bot. Gard. 32(1): 6-11.

Morton, C.V. 1962. A re-examination of Mexican Smilax (Smilacaceae). Brittonia 14: 299-309.

Schulz, O.E. 1904-1908. Smilax. Pp 17-47. In: I. Urban (ed.), Symbolae Antillanae seu Fundamenta Florae Indiae Occidentalis, vol. 5. Paul Klincksieck, Paris; Williams & Norgate, London.

Sipman, H. 1979. Liliaceae. Pp. 442-456. In: A.L. Stoffers & J.C. Lindeman (eds.), Flora of Suriname, Vol. 5, Part 1. Foundation Van Eedenfonds, Amsterdam.

Smilacaceae, F.N. Hepper. Flora of West Tropical Africa 3:1. 1968

Morphology General Habit

Shrubs, climbing or straggling, often with tendril-like petioles and prickly stems and branches; roots from an often stout rhizome; stems leafy

Morphology Leaves

Leaves alternate or opposite, 3-nerved, reticulate-veiny between the nerves

Morphology Reproductive morphology Flowers

Flowers dioecious or rarely bisexual, small, arranged in axillary umbels, racemes or spikes

Morphology Reproductive morphology Flowers Perianth

Perianth-segments 6, free or rarely united

Morphology Reproductive morphology Flowers Androecium

Stamens 6; filaments free or united; anthers apparently 1-locular by the confluence of the cells, introrse

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, 3-locular; ovules 1-2 in each loculus, pendulous

Morphology Reproductive morphology Flowers Sterile Parts

Staminodes present in the female flower

Morphology Reproductive morphology Fruits

Fruit a berry

Morphology Reproductive morphology Seeds

Seeds 1-3; embryo small in hard endosperm