Ceratozamia is a genus of New World cycads in the family Zamiaceae. The genus contains 27 known currently living species and one or two fossil species. Most species are endemic to mountainous areas of Mexico, while few species extend into the mountains of Guatemala, Honduras and Belize.[2][3] The genus name comes from the Greek ceras, meaning horn, which refers to the paired, spreading horny projections on the male and female sporophylls of all species.[4][5]

Ceratozamia
Ceratozamia mexicana
CITES Appendix I (CITES)[1]
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Gymnospermae
Division: Cycadophyta
Class: Cycadopsida
Order: Cycadales
Family: Zamiaceae
Subfamily: Zamioideae
Tribe: Ceratozamieae
D.W.Stev.
Genus: Ceratozamia
Brongn.
Type species
Ceratozamia mexicana
Synonyms[2]
  • Dipsacozamia Lehm.
  • Eriozamia Schuster nom. nud.

Many species have extremely limited ranges, and almost all described species are listed as vulnerable, endangered, or critically endangered by the IUCN Red List. The whole genus is listed under CITES Appendix I / EU Annex A, which prohibits international trade in specimens of these species except when the purpose of the import is not commercial, for instance for scientific research. Illegal plant poaching has posed a major threat to Ceratozamia species.[6]

Description

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The plants are dioecious, with a globose or cylindrical stem, rarely dichotomously branched, that may be underground or emergent. Several species produce basal shoots or suckers. The leaves are pinnately compound, straight, and spirally arranged. Leaf bases are usually deciduous but sometimes persistent. The petioles and rachis often have spines, though there may be very few to none. Leaflets are simple, entire, and articulate at the base, with parallel side veins and no distinct central vein. Male cones are cylindrical, upright, hairy, and stalked. Female cones are stalked or sessile, erect, and have short hairs. Seeds are oblong or elliptical, with a fleshy whitish outer coat.

Distribution and habitat

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Most species inhabit mountainous areas at 800–1000 m elevation, on sheltered slopes in moist forests. These forests range from tropical rainforests that are always wet, to pine-oak forests with alternating wet and dry seasons. There is a noticeable correlation between characteristics of species and the wetness or dryness of the habitat. Species with broad, thin leaflets live in wet habitats, and species with narrow, thick leaflets live in climates with wet and dry seasons.

Classification

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The genus consists of 27 known species:[2][7]

Phylogeny of Ceratozamia[8][9]
Image Leaves Scientific name Distribution
Ceratozamia alvarezii Pérez-Farr., Vovides & Iglesias Chiapas
Ceratozamia aurantiaca Pérez-Farrera et al. Chiapas
Ceratozamia becerrae Pérez-Farr., Vovides & Schutzman Tabasco, Chiapas
Ceratozamia brevifrons Miq. Veracruz
Ceratozamia chamberlainii Martínez-Domínguez Chiapas
Ceratozamia chimalapensis Pérez-Farr. & Vovides Oaxaca
Ceratozamia decumbens Vovides, S.Avendaño, Pérez-Farr. & González-Astorga Veracruz
Ceratozamia delucana Vázquez-Torres, Moretti & Carvajal-Hernández 2009 ex Vázquez-Torres, Moretti & Carvajal-Hernández
Ceratozamia dominguezii Pérez-Farrera & Gutiérrez-Ortega
Ceratozamia euryphyllidia Vázq. Torres, Sabato & D.W. Stev. Veracruz, Oaxaca
Ceratozamia fuscoviridis D. Moore Hidalgo
    Ceratozamia hildae G.P. Landry & M.C. Wilson Querétaro, San Luis Potosí
Ceratozamia hondurensis J.L. Haynes, Whitelock, Schutzman & R.S. Adams Honduras
Ceratozamia huastecorum S. Avendaño, Vovides & Cast.-Campos Veracruz
    Ceratozamia kuesteriana Regel Tamaulipas
    Ceratozamia latifolia Miq. Querétaro, San Luis Potosí, Hidalgo
Ceratozamia leptoceras Martínez-Domínguez et al.
Ceratozamia matudae Lundell Chiapas, Guatemala
    Ceratozamia mexicana Brongn. Puebla, Veracruz
  Ceratozamia microstrobila Vovides & J.D. Rees San Luis Potosí
    Ceratozamia miqueliana H. Wendl. Veracruz, Tabasco, Chiapas
Ceratozamia mirandae Vovides, Pérez-Farr. & Iglesias Chiapas
Ceratozamia mixeorum Chemnick, T.J. Greg. & S. Salas-Mor. Oaxaca
Ceratozamia morettii Vázq. Torres & Vovides Veracruz
    Ceratozamia norstogii D.W. Stev. Oaxaca, Chiapas
Ceratozamia oliversacksii Stevenson, Martínez-Domínguez & Nicolalde-Morejón
Ceratozamia osbornei Stevenson, Martínez-Domínguez & Nicolalde-Morejón
Ceratozamia reesii Vovides, Pérez Farrera & Gutiérrez-Ortega
    Ceratozamia robusta Miq. Veracruz, Oaxaca, Chiapas, Belize, Guatemala
  Ceratozamia sabatoi Vovides, Vázq. Torres, Schutzman & Iglesias Querétaro, Hidalgo
Ceratozamia sancheziae Pérez-Farr., Gutiérrez-Ortega & Vovides
Ceratozamia santillanii Pérez-Farr. & Vovides Chiapas
Ceratozamia schiblii Pérez-Farrera & Gutiérrez-Ortega
Ceratozamia subroseophylla Martínez-Domínguez & Nicolalde-Morejón
Ceratozamia tenuis (Dyer 1884) Stevenson & Vovides
Ceratozamia totonacorum Martínez-Domínguez & Nicolalde-Morejón
Ceratozamia vovidesii Pérez-Farr. & Iglesias Chiapas
Ceratozamia whitelockiana Chemnick & T.J. Greg. Oaxaca
  Ceratozamia zaragozae Medellin-Leal San Luis Potosí
Ceratozamia zoquorum Pérez-Farr., Vovides & Iglesias Chiapas

Fossil record

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There are several described fossil species, among them †Ceratozamia hofmannii and †Ceratozamia wrightii. Ceratozamia wrightii is the first evidence of the genus in the fossil record, with leaf fragments of the species found in Eocene deposits on Kupreanof Island in Alaska. This would support the hypothesis that there was a subtropical climate in northern areas during the Tertiary.

A fossil leaflet fragment of †Ceratozamia floersheimensis from the Rupelian stage of the Lower Oligocene has been found in marine sediments of the Bodenheim Formation in Rauenberg, Baden-Württemberg, Germany. Apart from Rauenberg and Flörsheim in Germany it is also known from the Oligocene of Budapest, Hungary and Trbovlje, Slovenia. A modern relationship exists to C. microstrobila, C. moretti, C. latifoli and C. delucana.[10]

A fossil leaf fragment of †Ceratozamia hofmannii has been recorded and described from Münzenberg near Leoben (upper Lower Miocene of Styria, Austria.[11] A fragmentary leaflet assigned to †C. hofmannii was recovered in the uppermost part of the Most Formation (Most Basin)) in North Bohemia, Czech Republic and dated by magnetostratigraphy and cyclostratigraphy to the last part of the early Miocene.[12]

References

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  1. ^ "Appendices | CITES". cites.org. Retrieved 2022-01-14.
  2. ^ a b c "World Checklist of Selected Plant Families: Royal Botanic Gardens, Kew". kew.org.
  3. ^ Hill, K.D. & Stevenson, D.W. (1999). A world list of Cycads, 1999. Excelsa 19: 67-72.
  4. ^ Standley, P. C. & J. A. Steyermark. 1958. Cycadaceae. In Standley, P.C. & Steyermark, J.A. (Eds), Flora of Guatemala – Part I. Fieldiana, Botany 24(1): 11–20.
  5. ^ Vovides, A. P., J. D. Rees & M. Vázquez-Torres. 1983. Zamiaceae. Flora de Veracruz 26: 1–31.
  6. ^ Christenhusz, M. J. M., J. L. Reveal, A. K. Farjon, M. F. Gardner, R. R. Mill & M. W. Chase. 2011. A new classification and linear sequence of extant gymnosperms. Phytotaxa 19: 55–70.
  7. ^ Haynes, J.L. 2011. World List of Cycads: A Historical Review Archived 2012-04-26 at the Wayback Machine. IUCN/SSC Cycad Specialist Group.
  8. ^ Stull, Gregory W.; Qu, Xiao-Jian; Parins-Fukuchi, Caroline; Yang, Ying-Ying; Yang, Jun-Bo; Yang, Zhi-Yun; Hu, Yi; Ma, Hong; Soltis, Pamela S.; Soltis, Douglas E.; Li, De-Zhu; Smith, Stephen A.; Yi, Ting-Shuang; et al. (2021). "Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms". Nature Plants. 7 (8): 1015–1025. bioRxiv 10.1101/2021.03.13.435279. doi:10.1038/s41477-021-00964-4. PMID 34282286. S2CID 232282918.
  9. ^ Stull, Gregory W.; et al. (2021). "main.dated.supermatrix.tree.T9.tre". Figshare. doi:10.6084/m9.figshare.14547354.v1. {{cite journal}}: Cite journal requires |journal= (help)
  10. ^ Early Oligocene plant diversity along the Upper Rhine, Graben: The fossil flora of Rauenberg, Germany by Johanna Kovar-Eder - Acta Palaeobotanica 56(2): 329-440, 2016 DOI: 10.1515/acpa-2016-0011
  11. ^ A noteworthy cycad, Ceratozamia hofmannii ETTINGSHAUSEN 1887, from the Lower Miocene of Austria re-examined by Zlatko Kvaček in Neues Jahrbuch fur Geologie und Palaontologie Monatshefte · February 2004
  12. ^ New fossil records of Ceratozamia (Zamiaceae, Cycadales) from the European Oligocene and lower Miocene by Zlatko Kvaček Acta Palaeobotanica 54(2):231-247 · December 2014

Bibliography

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  • Jones, David L. (2002). Cycads of the World: Ancient Plants in Today's Landscape. Washington, D.C.: Smithsonian Institution Press. ISBN 1-58834-043-0.
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