Ceratozamia
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''Ceratozamia'' is a genus of New World
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
s in the family
Zamiaceae The Zamiaceae are a family of cycads that are superficially palm or fern-like. They are divided into two subfamilies with eight genera and about 150 species in the tropical and subtropical regions of Africa, Australia and North and South America. ...
. The genus contains 27 known currently living species and one or two fossil species. Most species are
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
to mountainous areas of
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
, while few species extend into the mountains of
Guatemala Guatemala ( ; ), officially the Republic of Guatemala ( es, República de Guatemala, links=no), is a country in Central America. It is bordered to the north and west by Mexico; to the northeast by Belize and the Caribbean; to the east by H ...
,
Honduras Honduras, officially the Republic of Honduras, is a country in Central America. The republic of Honduras is bordered to the west by Guatemala, to the southwest by El Salvador, to the southeast by Nicaragua, to the south by the Pacific Oce ...
and
Belize Belize (; bzj, Bileez) is a Caribbean and Central American country on the northeastern coast of Central America. It is bordered by Mexico to the north, the Caribbean Sea to the east, and Guatemala to the west and south. It also shares a wate ...
. The genus name comes from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''ceras'', meaning horn, which refers to the paired, spreading horny projections on the male and female
sporophyll A sporophyll is a leaf that bears sporangia. Both microphylls and megaphylls can be sporophylls. In heterosporous plants, sporophylls (whether they are microphylls or megaphylls) bear either megasporangia and thus are called megasporophylls, or mi ...
s of all species. 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 International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
. The whole genus is listed under
CITES Appendix I CITES (shorter name for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of interna ...
/ 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 Plant collecting is the acquisition of plant specimens for the purposes of research, cultivation, or as a hobby. Plant specimens may be kept alive, but are more commonly dried and pressed to preserve the quality of the specimen. Plant collecting i ...
has posed a major threat to ''Ceratozamia'' species.


Description

The plants are
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
, with a globose or cylindrical stem, rarely dichotomously branched, that may be underground or emergent. Several species produce
basal shoot Basal shoots, root sprouts, adventitious shoots, and suckers are words for various kinds of shoots that grow from adventitious buds on the base of a tree or shrub, or from adventitious buds on its roots. Shoots that grow from buds on the base of ...
s or suckers. The
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
are pinnately compound, straight, and spirally arranged. Leaf bases are usually deciduous but sometimes persistent. The petioles and
rachis In biology, a rachis (from the grc, ῥάχις [], "backbone, spine") is a main axis or "shaft". In zoology and microbiology In vertebrates, ''rachis'' can refer to the series of articulated vertebrae, which encase the spinal cord. In this c ...
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

Most species inhabit mountainous areas at 800–1000 m elevation, on sheltered slopes in moist forests. These forests range from
tropical rainforest Tropical rainforests are rainforests that occur in areas of tropical rainforest climate in which there is no dry season – all months have an average precipitation of at least 60 mm – and may also be referred to as ''lowland equatori ...
s 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

The genus consists of 27 known species:


Fossil record

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 The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
deposits on
Kupreanof Island Kupreanof Island (russian: Остров Купреянова) is an island in the Alexander Archipelago in southeastern Alaska. The island is long and wide with a total land area is , making it the 13th largest island in the United States and t ...
in
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S., ...
. This would support the hypothesis that there was a subtropical climate in northern areas during the
Tertiary Tertiary ( ) is a widely used but obsolete term for the geologic period from 66 million to 2.6 million years ago. The period began with the demise of the non-avian dinosaurs in the Cretaceous–Paleogene extinction event, at the start ...
. A
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
leaflet fragment of †''Ceratozamia floersheimensis'' from the
Rupelian stage The Rupelian is, in the geologic timescale, the older of two ages or the lower of two stages of the Oligocene Epoch/Series. It spans the time between . It is preceded by the Priabonian Stage (part of the Eocene) and is followed by the Chattian S ...
of the
Lower Oligocene The Rupelian is, in the geologic timescale, the older of two ages or the lower of two stages of the Oligocene Epoch/Series. It spans the time between . It is preceded by the Priabonian Stage (part of the Eocene) and is followed by the Chatti ...
has been found in
marine sediments Marine sediment, or ocean sediment, or seafloor sediment, are deposits of insoluble particles that have accumulated on the seafloor. These particles have their origins in soil and rocks and have been transported from the land to the sea, mainly ...
of the
Bodenheim Bodenheim is a state-recognized tourism municipality (''Fremdenverkehrsgemeinde'') in the Mainz-Bingen district in Rhineland-Palatinate, Germany Geography Location The municipality lies roughly 12 km south of downtown Mainz on the edge of t ...
Formation in Rauenberg,
Baden-Württemberg Baden-Württemberg (; ), commonly shortened to BW or BaWü, is a German state () in Southwest Germany, east of the Rhine, which forms the southern part of Germany's western border with France. With more than 11.07 million inhabitants across a ...
,
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. Apart from Rauenberg and Flörsheim in
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
it is also known from the
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
of
Budapest Budapest (, ; ) is the capital and most populous city of Hungary. It is the ninth-largest city in the European Union by population within city limits and the second-largest city on the Danube river; the city has an estimated population ...
,
Hungary Hungary ( hu, Magyarország ) is a landlocked country in Central Europe. Spanning of the Carpathian Basin, it is bordered by Slovakia to the north, Ukraine to the northeast, Romania to the east and southeast, Serbia to the south, Croatia a ...
and
Trbovlje Trbovlje (; german: Trifail''Leksikon občin kraljestev in dežel zastopanih v državnem zboru,'' vol. 4: ''Štajersko''. 1904. Vienna: C. Kr. Dvorna in Državna Tiskarna, p. 58.) is Slovenia's tenth-largest town, and the seat of the Municipality o ...
,
Slovenia Slovenia ( ; sl, Slovenija ), officially the Republic of Slovenia (Slovene: , abbr.: ''RS''), is a country in Central Europe. It is bordered by Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the southeast, an ...
. A modern relationship exists to ''C. microstrobila'', ''C. moretti'', ''C. latifoli'' and ''C. delucana''. A
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
leaf fragment of †''Ceratozamia hofmannii'' has been recorded and described from Münzenberg near
Leoben Leoben () is a Styrian city in central Austria, located on the Mur river. With a population of about 25,000 it is a local industrial centre and hosts the University of Leoben, which specialises in mining. The Peace of Leoben, an armistice betwee ...
(upper
Lower Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene Epoch made up of two stages: the Aquitanian and Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 Ma to 15.97 ± 0.05 Ma (million years ago). It was prece ...
of
Styria Styria (german: Steiermark ; Serbo-Croatian and sl, ; hu, Stájerország) is a state (''Bundesland'') in the southeast of Austria. With an area of , Styria is the second largest state of Austria, after Lower Austria. Styria is bordered to ...
,
Austria Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...
. A fragmentary leaflet assigned to †''C. hofmannii'' was recovered in the uppermost part of the Most Formation (
Most Basin The Most Basin (also known as North Bohemian Basin; cs, Mostecká pánev, german: Nordböhmisches Becken) is a tectonic Depression (geology), depression and Geomorphological division of the Czech Republic, geomorphological mesoregion of the Czech ...
)) in
North Bohemia North Bohemia ( cs, Severní Čechy, german: Nordböhmen) is a region in the north of the Czech Republic. Location North Bohemia roughly covers the present-day NUTS regional unit of ''CZ04 Severozápad'' and the western part of ''CZ05 Severovýc ...
,
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
and dated by
magnetostratigraphy Magnetostratigraphy is a geophysical correlation technique used to date sedimentary and volcanic sequences. The method works by collecting oriented samples at measured intervals throughout the section. The samples are analyzed to determine their '' ...
and
cyclostratigraphy Cyclostratigraphy is a subdiscipline of stratigraphy that studies astronomically forced climate cycles within sedimentary successions. Orbital changes Astronomical cycles (also known as Milankovitch cycles) are variations of the Earth's or ...
to the last part of the early
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
.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


References


Bibliography

*


External links


''Ceratozamia'' at The Cycad Pages


{{Taxonbar, from=Q135250 Zamiaceae Taxa named by Adolphe-Théodore Brongniart