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Cucurbitaceae

Cucurbitaceae
Temporal range: Early Paleocene to Recent, 62–0 Ma
Hodgsonia heteroclita male.jpg
Hodgsonia male plant
Scientific classification e
Kingdom: Plantae
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Cucurbitales
Family: Cucurbitaceae
Juss.[1]
Type genus
Cucurbita
Tribes and genera

See text.

The Cucurbitaceae, also called cucurbits or the gourd family, are a plant family consisting of about 965 species in around 95 genera,[2] the most important of which are:

The plants in this family are grown around the tropics and in temperate areas, where those with edible fruits were among the earliest cultivated plants in both the Old and New Worlds. The Cucurbitaceae family ranks among the highest of plant families for number and percentage of species used as human food.[3] The name Cucurbitaceae comes to international scientific vocabulary from New Latin, from Cucurbita, the type genus, + -aceae,[4] a standardized suffix for plant family names in modern taxonomy. The genus name comes from the Classical Latin word cucurbita, "gourd".

Description

Most of the plants in this family are annual vines, but some are woody lianas, thorny shrubs, or trees (Dendrosicyos). Many species have large, yellow or white flowers. The stems are hairy and pentangular. Tendrils are present at 90° to the leaf petioles at nodes. Leaves are exstipulate alternate simple palmately lobed or palmately compound. The flowers are unisexual, with male and female flowers on different plants (dioecious) or on the same plant (monoecious). The female flowers have inferior ovaries. The fruit is often a kind of modified berry called a pepo.

Fossil history

One of the oldest fossil cucurbits so far is †Cucurbitaciphyllum lobatum from the Paleocene epoch, found at Shirley Canal, Montana. It was described for the first time in 1924 by the paleobotanist Frank Hall Knowlton. The fossil leaf is palmate, trilobed with rounded lobal sinuses and an entire or serrate margin. It has a leaf pattern similar to the members of the genera Kedrostis, Melothria and Zehneria.[5]

Classification

Tribal classification

Pumpkins and squashes displayed in a show competition
A selection of cucurbits of the South Korean Genebank in Suwon
Cucurbits on display at the Real Jardín Botánico de Madrid, with the title "Variedades de calabaza"

The most recent classification of Cucurbitaceae delineates 15 tribes:[6][7]

  • Tribe Gomphogyneae Benth. & Hook. f. 1867
    • Alsomitra (Blume) Spach 1838 (1 sp.)
    • Bayabusua (1 sp.)
    • Gomphogyne Griff. 1845 (2 spp.)
    • Gynostemma Blume 1825 (10 spp.)
    • Hemsleya Cogn. ex F.B. Forbes & Hemsl. 1888 (30 spp.)
    • Neoalsomitra Hutch. 1942 (12 spp.)
  • Tribe Triceratieae A. Rich. 1845
    • Anisosperma Silva Manso 1836 (1 sp.)
    • Cyclantheropsis Harms 1896 (3 spp.)
    • Fevillea L. 1753 (8 spp.)
    • Pteropepon (Cogn.) Cogn. 1916 (5 spp.)
    • Sicydium Schltdl. 1832 (7 spp.)
  • Tribe Zanonieae Benth. & Hook. f. 1867
  • Tribe Actinostemmateae H. Schaef. & S.S. Renner 2011
    • Actinostemma Griff. 1841 (3 spp.)
  • Tribe Indofevilleeae H. Schaef. & S.S. Renner 2011
    • Indofevillea Chatterjee 1946 (1 sp.)
  • Tribe Thladiantheae H. Schaef. & S.S. Renner 2011
    • Baijiania A.M. Lu & J.Q. Li 1993 (30 spp.)
    • Thladiantha Bunge 1833 (5 spp.)
  • Tribe Siraitieae H. Schaef. & S.S. Renner 2011
  • Tribe Momordiceae H. Schaef. & S.S. Renner 2011
  • Tribe Joliffieae Schrad. 1838
    • Ampelosicyos Thouars 1808 (5 spp.)
    • Cogniauxia Baill. 1844 (2 spp.)
    • Telfairia Hook. 1827 (3 spp.)
  • Tribe Bryonieae Dumort. 1827
    • Austrobryonia H. Schaef. 2008 (4 spp.)
    • Bryonia L. 1753 (10 spp.)
    • Ecballium A. Rich. 1824 (1 sp.)
  • Tribe Schizopeponeae C. Jeffrey 1964
    • Herpetospermum Wall. ex Hook. f. 1867 (3 spp.)
    • Schizopepon Maxim. 1859 (6–8 spp.)
  • Tribe Sicyoeae Schrad. 1838
  • Tribe Coniandreae Endl. 1839
    • Apodanthera Arn. 1841 (16 spp.)
    • Bambekea Cogn. 1916 (1 sp.)
    • Ceratosanthes Adans. 1763 (4 spp.)
    • Corallocarpus Welw. ex Benth. & Hook. f. 1867 (17 spp.)
    • Cucurbitella Walp. 1846 (1 sp.)
    • Dendrosicyos Balf. f. 1882 (1 sp.)
    • Doyerea Grosourdy 1864 (1 sp.)
    • Eureiandra Hook. f. 1867 (8 spp.)
    • Gurania (Schltdl.) Cogn. 1875 (37 spp.)
    • Halosicyos Mart. Crov 1947 (1 sp.)
    • Helmontia Cogn. 1875 (2–4 spp.)
    • Ibervillea Greene 1895 (9–10 spp.)
    • Kedrostis Medik. 1791 (28 spp.)
    • Melotrianthus M. Crovetto 1954 (1–3 spp.)
    • Psiguria Neck. ex Arn. 1841 (6–12 spp.)
    • Seyrigia Keraudren 1960 [1961] (6 spp.)
    • Trochomeriopsis Cogn. 1881 (1 sp.)
    • Tumamoca Rose 1912 (2 spp.)
    • Wilbrandia Silva Manso 1836 (5 spp.)
  • Tribe Benincaseae Ser. 1825
    • Acanthosicyos Welw. ex Hook. f. 1867 (1 sp.)
    • Benincasa Savi 1818 (2 spp., including Praecitrullus Pangalo 1944)
    • Borneosicyos (1–2 spp.)
    • Cephalopentandra Chiov. 1929 (1 sp.)
    • Citrullus Schrad. 1836 (4 spp.)
    • Coccinia Wight & Arn. 1834 (30 spp.)
    • Ctenolepis Hook. f. 1867 (3 spp.)
    • Cucumis L. 1753 (65 spp.)
    • Dactyliandra Hook. f. 1871 (2 spp.)
    • Diplocyclos (Endl.) T. Post & Kuntze 1903 (4 spp.)
    • Indomelothria (2 spp.)
    • Khmeriosicyos (1 sp.)
    • Lagenaria Ser. 1825 (6 spp.)
    • Lemurosicyos Keraudren 1963 [1964] (1 sp.)
    • Melothria L. 1753 (12 spp.)
    • Muellerargia Cogn. 1881 (2 sp.)
    • Papuasicyos (8 spp.)
    • Peponium Engl. 1897 (20 spp.)
    • Raphidiocystis Hook. f. 1867 (5 spp.)
    • Ruthalicia C. Jeffrey 1962 (2 spp.)
    • Scopellaria W.J. de Wilde & Duyfjes 2006 (2 spp.)
    • Solena Lour. 1790 (3 spp.)
    • Trochomeria Hook. f. 1867 (8 spp.)
    • Zehneria Endl. 1833 (ca. 60 spp.)

Systematics

Modern molecular phylogenetics suggest the following relationships:[6][8][9][10][11][12]

Anisophylleaceae (outgroup)

Cucurbitaceae
Gomphogyneae

Alsomitra

Bayabusua

Hemsleya

Gomphogyne

Gynostemma

Neoalsomitra

Triceratieae

Fevillea

Pteropepon

Cyclantheropsis

Sicydium

Zanonieae

Gerrardanthus

Zanonia

Siolmatra

Xerosicyos

Actinostemmateae

Actinostemma

Indofevilleeae

Indofevillea

Thladiantheae

Baijiania

Thladiantha

Siraitieae

Siraitia

Momordiceae

Momordica

Joliffieae

Cogniauxia

Telfairia

Ampelosicyos

Bryonieae

Ecballium

Bryonia

Austrobryonia

Sicyoeae

Nothoalsomitra

Hodgsonia

Echinocystis

Marah

Echinopepon

Frantzia

Cyclanthera

Hanburia

Sicyos

Linnaeosicyos

Luffa

Trichosanthes

Schizopeponeae

Schizopepon

Herpetospermum

Coniandreae

Bambekea

Eureiandra

Dendrosicyos

Trochomeriopsis

Seyrigia

Corallocarpus

Cucurbitella

Doyerea

{{Expansion depth limit exceeded|1=Wilbrandia

1=Psiguria 1=Helmontia 2=Gurania
                                            }}
                                          }}
                                        }}

Melotrianthus

Kedrostis

{{Expansion depth limit exceeded|1={{Expansion depth limit exceeded|1={{Expansion depth limit exceeded|1=Ceratosanthes

2=Halosicyos
                                            }}
2=Apodanthera
                                          }}
1=Tumamoca 2=Ibervillea
                                          }}
                                        }}
Cucurbiteae

Polyclathra

Peponopsis

Cucurbita

Penelopeia

Calycophysum

Sicana

Tecunumania

Schizocarpum

Cionosicys

Abobra

Cayaponia

Benincaseae

Zehneria

Citrullus

Peponium

Lagenaria

Acanthosicyos

Raphidiocystis

{{Expansion depth limit exceeded|1=Cephalopentandra

1=Lemurosicyos 1=Solena 2=Borneosicyos
                                            }}
                                          }}
                                        }}

Benincasa

{{Expansion depth limit exceeded|1=Ctenolepis

1=Dactyliandra 2=Trochomeria
                                          }}
                                        }}

Ruthalicia

Indomelothria

Melothria

Coccinia

Diplocyclos

Papuasicyos

Scopellaria

Muellerargia

Cucumis

References

  1. ^ Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x.
  2. ^ Christenhusz, M. J. M. & Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3): 201–217. doi:10.11646/phytotaxa.261.3.1.
  3. ^ "Cucurbits". Purdue University. Retrieved August 26, 2013.
  4. ^ Merriam-Webster, Merriam-Webster's Unabridged Dictionary, Merriam-Webster.
  5. ^ Revisions to Roland Brown's North American Paleocene Flora by Steven R. Manchester at Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA. Published in Acta Musei Nationalis Pragae, Series B – Historia Naturalis, vol. 70, 2014, no. 3-4, pp. 153–210.
  6. ^ a b Schaefer H, Renner SS (2011). "Phylogenetic relationships in the order Cucurbitales and a new classification of the gourd family (Cucurbitaceae)". Taxon. 60 (1): 122–138. doi:10.1002/tax.601011. JSTOR 41059827. Archived from the original on 31 January 2018. Retrieved 20 March 2017.
  7. ^ Schaefer H, Kocyan A, Renner SS (2007). "Phylogenetics of Cucumis (Cucurbitaceae): Cucumber (C. sativus) belongs in an Asian/Australian clade far from melon (C. melo)". BMC Evolutionary Biology. 7: 58–69. doi:10.1186/1471-2148-7-58. PMC 3225884. PMID 17425784.
  8. ^ Zhang L-B, Simmons MP, Kocyan A, Renner SS (2006). "Phylogeny of the Cucurbitales based on DNA sequences of nine loci from three genomes: Implications for morphological and sexual system evolution" (PDF). Molecular Phylogenetics and Evolution. 39 (2): 305–322. doi:10.1016/j.ympev.2005.10.002. PMID 16293423.
  9. ^ Schaefer H, Heibl C, Renner SS (2009). "Gourds afloat: A dated phylogeny reveals an Asian origin of the gourd family (Cucurbitaceae) and numerous oversea dispersal events". Proc Royal Soc B. 276 (1658): 843–851. doi:10.1098/rspb.2008.1447. PMC 2664369. PMID 19033142.
  10. ^ de Boer HJ, Schaefer H, Thulin M, Renner SS (2012). "Evolution and loss of long-fringed petals: A case study using a dated phylogeny of the snake gourds, Trichosanthes (Cucurbitaceae)". BMC Evolutionary Biology. 12: 108. doi:10.1186/1471-2148-12-108. PMC 3502538. PMID 22759528.
  11. ^ Belgrano MJ (2012). Estudio sistemático y biogeográfico del género Apodanthera Arn. (Cucurbitaceae) [Systematic and biogeographic study of the genus Apodanthera Arn. (Cucurbitaceae)] (Ph.D.). Universidad Nacional de La Plata.
  12. ^ Renner SS, Schaefer H (2016). "Phylogeny and evolution of the Cucurbitaceae". In Grumet R, Katzir N, Garcia-Mas J (eds.). Genetics and Genomics of Cucurbitaceae. Plant Genetics and Genomics: Crops and Models. 20. New York, NY: Springer International Publishing. pp. 1–11. doi:10.1007/7397_2016_14. ISBN 978-3-319-49330-5.

Further reading

  • Bates D, Robinson R, Jeffrey C, eds. (1990). Biology and Utilization of the Cucurbitaceae. Cornell University Press. ISBN 978-0-8014-1670-5.
  • Jeffrey C. (2005). "A new system of Cucurbitaceae". Bot. Zhurn. 90: 332–335.

External links