Cucurbitaceae

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

See text.

The Cucurbitaceae (/kjuːˌkɜːrbɪˈteɪsii/), also called cucurbits and 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 Cucurbitaceae consist of 98 proposed genera with 975 species,[4] mainly in regions tropical and subtropical. All species are sensitive to frost. 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.)

Alphabetical list of genera

Abobra, Acanthosicyos, Actinostemma, Alsomitra, Ampelosicyos, Anisosperma, Apodanthera, Austrobryonia, Baijiania, Bambekea, Bayabusua, Benincasa, Borneosicyos, Bryonia, Calycophysum, Cayaponia, Cephalopentandra, Ceratosanthes, Cionosicys, Citrullus, Coccinia, Cogniauxia, Corallocarpus, Ctenolepis, Cucumis, Cucurbita, Cucurbitella, Cyclanthera, Cyclantheropsis, Dactyliandra, Dendrosicyos, Diplocyclos, Doyerea, Ecballium, Echinocystis, Echinopepon, Eureiandra, Fevillea, Frantzia, Gerrardanthus, Gomphogyne, Gurania, Gynostemma, Halosicyos, Hanburia, Helmontia, Hemsleya, Herpetospermum, Hodgsonia, Ibervillea, Indofevillea, Indomelothria, Kedrostis, Khmeriosicyos, Lagenaria, Lemurosicyos, Linnaeosicyos, Luffa, Marah, Melothria, Melotrianthus, Momordica, Muellerargia, Neoalsomitra, Nothoalsomitra, Papuasicyos, Penelopeia, Peponium, Peponopsis, Polyclathra, Psiguria, Pteropepon, Raphidiocystis, Ruthalicia, Schizocarpum, Schizopepon, Scopellaria, Seyrigia, Sicana, Sicydium, Sicyos, Siolmatra, Siraitia, Solena, Tecunumania, Telfairia, Thladiantha, Trichosanthes, Trochomeria, Trochomeriopsis, Tumamoca, Wilbrandia, Xerosicyos, Zanonia, Zehneria.

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

 
 

 

Wilbrandia

 
 

 

Psiguria

 
 

 

Helmontia

 
 

Gurania

 
 
 
 
 
 

Melotrianthus

 
 

 

Kedrostis

 
 

 

 

 

Ceratosanthes

 
 

Halosicyos

 
 
 

Apodanthera

 
 
 

 

Tumamoca

 
 

Ibervillea

 
 
 
 
 
 
 
 
 
 
 

Cucurbiteae

 

 

Polyclathra

 
 

 

Peponopsis

 
 

Cucurbita

 
 
 
 

 

 

Penelopeia

 
 

Calycophysum

 
 

Sicana

 
 
 

 

Tecunumania

 
 

 

Schizocarpum

 
 

 

Cionosicys

 
 

 

Abobra

 
 

Cayaponia

 
 
 
 
 
 
 
Benincaseae

 

Zehneria

 
 

 

 

 

Citrullus

 
 

 

Peponium

 
 

Lagenaria

 
 
 
 

 

Acanthosicyos

 
 

 

 

Raphidiocystis

 
 

 

Cephalopentandra

 
 

 

Lemurosicyos

 
 

 

Solena

 
 

Borneosicyos

 
 
 
 
 
 

 

Benincasa

 
 

 

Ctenolepis

 
 

 

Dactyliandra

 
 

Trochomeria

 
 
 
 
 
 
 
 

 

 

Ruthalicia

 
 

 

Indomelothria

 
 

Melothria

 
 
 
 

 

 

 

Coccinia

 
 

Diplocyclos

 
 
 

 

Papuasicyos

 
 

Scopellaria

 
 
 
 

 

Muellerargia

 
 

Cucumis

 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 

Images of cucurbits in Byzantine mosaics from Israel

Round melons and elongated adzhur melons in Kursi church mosaic, Israel, near the Sea of Galilee

Six cucurbit crops are represented in 23 Byzantine-era mosaics from Israel, these being round melons (Cucumis melo), watermelons (Citrullus lanatus), sponge gourds (Luffa aegyptiaca), snake melons (faqqous, Cucumis melo flexuosus group), adzhur melons (C. melo adzhur group), and bottle gourds (Lagenaria siceraria). Cucurbits are represented in 23 of the 134 mosaics containing images of crop plants, a surprisingly high frequency of 17%. Several of the cucurbit images have not been found elsewhere, suggesting a diverse and highly developed local horticulture of cucurbits in Israel during the Byzantine era. Representations of mature sponge gourds are found in widespread localities, suggestive of the high value accorded to cleanliness and hygiene.[13]

Etymology and pronunciation

The name Cucurbitaceae comes to international scientific vocabulary from New Latin, from Cucurbita, the type genus, + -aceae,[14] a standardized suffix for plant family names in modern taxonomy. The genus name comes from the Classical Latin word cucurbita, "gourd".

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. ↑ "Angiosperm Phylogeny Website". mobot.org.
  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. 1 2 Schaefer H, Renner SS (2011). "Phylogenetic relationships in the order Cucurbitales and a new classification of the gourd family (Cucurbitaceae)" (PDF). Taxon. 60 (1): 122–138. JSTOR 41059827. Archived from the original on 27 February 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. Archived from the original on 27 February 2017.
  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" (PDF). Proc Royal Soc B. 276 (1658): 843–851. doi:10.1098/rspb.2008.1447. PMC 2664369. PMID 19033142. Archived from the original on 27 February 2017.
  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. 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.
  13. ↑ Avital, Anat; Paris, Harry S. (2014-08-01). "Cucurbits depicted in Byzantine mosaics from Israel, 350–600 ce". Annals of Botany. 114 (2): 203–222. doi:10.1093/aob/mcu106. ISSN 0305-7364. PMC 4111391. PMID 24948671.
  14. ↑ Merriam-Webster, Merriam-Webster's Unabridged Dictionary, Merriam-Webster.

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.
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