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<document ID-DOI="http://dx.doi.org/10.3897/phytokeys.100.23152" ID-PMC="PMC6023955" ID-Pensoft-Pub="1314-2003-100-91" ID-Pensoft-UUID="117AFFB5FFE90945FB26EE02FF92FFA4" ID-PubMed="29962891" ID-Zenodo-Dep="1300062" ModsDocID="1314-2003-100-91" checkinTime="1529641986672" checkinUser="pensoft" docAuthor="Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco &amp; Stevenson, Dennis Wm." docDate="2018" docId="68E6B809971D5A51BD203414DBAC2F83" docLanguage="en" docName="PhytoKeys 100: 91-124" docOrigin="PhytoKeys 100" docPubDate="2018-06-21" docSource="http://dx.doi.org/10.3897/phytokeys.100.23152" docTitle="Ceratozamia mexicana Brongn. Ann. Sci. Nat., Bot. ser. 3, 5: 7 8, t. 1. 1846." docType="treatment" docVersion="3" id="117AFFB5FFE90945FB26EE02FF92FFA4" lastPageNumber="107" masterDocId="117AFFB5FFE90945FB26EE02FF92FFA4" masterDocTitle="Taxonomic review of Ceratozamia (Zamiaceae) in the Sierra Madre Oriental, Mexico" masterLastPageNumber="124" masterPageNumber="91" pageNumber="105" updateTime="1668138916235" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Taxonomic review of Ceratozamia (Zamiaceae) in the Sierra Madre Oriental, Mexico</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Martinez-Dominguez, Lili</mods:namePart>
<mods:affiliation>Laboratorio de Taxonomia Integrativa, Instituto de Investigaciones Biologicas, Universidad Veracruzana, Xalapa, 91190, Veracruz, Mexico &amp; Centro de Investigaciones Tropicales, Universidad Veracruzana, Jose Maria Morelos 44, Zona Centro, Xalapa, 91000, Veracruz, Mexico</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Nicolalde-Morejon, Fernando</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0003-1423-7474</mods:nameIdentifier>
<mods:affiliation>Laboratorio de Taxonomia Integrativa, Instituto de Investigaciones Biologicas, Universidad Veracruzana, Xalapa, 91190, Veracruz, Mexico</mods:affiliation>
<mods:nameIdentifier type="email">f_nicolalde@yahoo.com</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Vergara-Silva, Francisco</mods:namePart>
<mods:affiliation>Laboratorio de Sistematica Molecular (Jardin Botanico), Instituto de Biologia, Universidad Nacional Autonoma de Mexico, 3 er Circuito Exterior, Ciudad Universitaria, Coyoacan 04510, Mexico, D. F., Mexico</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Stevenson, Dennis Wm.</mods:namePart>
<mods:affiliation>The New York Botanical Garden, Bronx, Nueva York, 10458 - 5120, USA</mods:affiliation>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>PhytoKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2018</mods:date>
<mods:detail type="pubDate">
<mods:number>2018-06-21</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>100</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>91</mods:start>
<mods:end>124</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location>
<mods:url>http://dx.doi.org/10.3897/phytokeys.100.23152</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/phytokeys.100.23152</mods:identifier>
<mods:identifier type="Pensoft-Pub">1314-2003-100-91</mods:identifier>
<mods:identifier type="Pensoft-UUID">117AFFB5FFE90945FB26EE02FF92FFA4</mods:identifier>
<mods:identifier type="Zenodo-Dep">1300062</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="182225063" LSID="urn:lsid:plazi:treatment:68E6B809971D5A51BD203414DBAC2F83" httpUri="http://treatment.plazi.org/id/68E6B809971D5A51BD203414DBAC2F83" lastPageId="16" lastPageNumber="107" pageId="14" pageNumber="105">
<subSubSection pageId="14" pageNumber="105" type="nomenclature">
<paragraph pageId="14" pageNumber="105">
9.
<taxonomicName LSID="68E6B809-971D-5A51-BD20-3414DBAC2F83" authority="Brongn. Ann. Sci. Nat., Bot. ser. 3, 5: 7 - 8, t. 1. 1846." authorityName="Brongn. Ann. Sci. Nat., Bot. ser. 3, 5: 7 8, t. 1. 1846." class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia mexicana" order="Cycadales" pageId="14" pageNumber="105" phylum="Tracheophyta" rank="species" species="mexicana">Ceratozamia mexicana Brongn. Ann. Sci. Nat., Bot. ser. 3, 5: 7-8, t. 1. 1846.</taxonomicName>
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Shapes of leaflets. A C. latifolia B C. kuesteriana C C. morettii D C. sabatoi E Ceratozamia brevifrons F C. chamberlainii G C. decumbens H C. fuscoviridis I C. tenuis J C. totonacorum K C. delucana L C. mexicana." figureDoi="10.3897/phytokeys.100.23152.figure3" httpUri="https://binary.pensoft.net/fig/211374" pageId="14" pageNumber="105">Figures 3L</figureCitation>
<figureCitation captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Ceratozamia mexicana. A Adult plant in habit B Leaf at emergence C Detail of pollen strobili D Microsporophyll E Megasporophyll F Detail of ovulate strobili G Seeds." figureDoi="10.3897/phytokeys.100.23152.figure11" httpUri="https://binary.pensoft.net/fig/211382" pageId="14" pageNumber="105">, 11</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="14" pageNumber="105" type="materials_examined">
<paragraph pageId="14" pageNumber="105">
<typeStatus>Type</typeStatus>
.
</paragraph>
<paragraph pageId="14" pageNumber="105">
<collectingCountry name="Mexico">MEXICO</collectingCountry>
, from Ghiesbrecht, cult. in Hort. Bot. Parisiensis, 1845,
<emphasis italics="true" pageId="14" pageNumber="105">Brongniart s.n.</emphasis>
(
<typeStatus>holotype</typeStatus>
: P). Epitype (designated by
<bibRefCitation DOI="https://doi.org/10.17129/botsci.449" author="Vovides, AP" journalOrPublisher="Botanical Journal of the Linnean Society" pageId="27" pageNumber="118" refId="B51" refString="Vovides, AP, Stevenson, D, Perez-Farrera, MA, Lopez, S, Avendano, S, 2016. What is Ceratozamia mexicana (Zamiaceae)? Botanical Sciences 94(2): 419-429. https://doi.org/10.17129/botsci.449" title="What is Ceratozamia mexicana (Zamiaceae)? Botanical Sciences 94 (2): 419 - 429." url="https://doi.org/10.17129/botsci.449" year="2016">Vovides et al. 2016</bibRefCitation>
).
<materialsCitation collectingDate="1982-09-08" collectorName="J. Rees, A. P. Vovides" country="MEXICO" location="Totutla" specimenCount="1" stateProvince="Veracruz" typeStatus="holotype">
<collectingCountry name="Mexico">MEXICO</collectingCountry>
.
<collectingRegion country="Mexico" name="Veracruz de Ignacio de la Llave">Veracruz</collectingRegion>
:
<location LSID="urn:lsid:plazi:treatment:68E6B809971D5A51BD203414DBAC2F83:EC31A88A0702478F007F7C5F0D9A4114" country="MEXICO" name="Totutla" stateProvince="Veracruz">Totutla</location>
,
<collectingDate value="1982-09-08">08 Sep. 1982</collectingDate>
,
<emphasis italics="true" pageId="14" pageNumber="105">
<collectorName>J. Rees</collectorName>
&amp;
<collectorName>A. P. Vovides</collectorName>
1688
</emphasis>
(XAL)
</materialsCitation>
.
</paragraph>
<paragraph pageId="14" pageNumber="105">
<taxonomicName authorityName="Miq" authorityYear="1847" class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia intermedia" order="Cycadales" pageId="14" pageNumber="105" phylum="Tracheophyta" rank="species" species="intermedia">
<emphasis italics="true" pageId="14" pageNumber="105">Ceratozamia intermedia</emphasis>
</taxonomicName>
<materialsCitation collectingDate="1985-03-13" collectorName="Miq. Tijdschr. Wis-Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch., G. Castillo-Campo, Medina" country="MEXICO" location="Puente Nacional" specimenCount="1" stateProvince="Veracruz" typeStatus="Neotype">
<collectorName>Miq. Tijdschr. Wis-Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch.</collectorName>
1 (4): 40-41. 1848.
<typeStatus>Neotype</typeStatus>
(designated here).
<collectingCountry name="Mexico">MEXICO</collectingCountry>
.
<collectingRegion country="Mexico" name="Veracruz de Ignacio de la Llave">Veracruz</collectingRegion>
:
<location LSID="urn:lsid:plazi:treatment:68E6B809971D5A51BD203414DBAC2F83:D3A7EB8B0D4874E2C7676F2FF37CB16B" country="MEXICO" name="Puente Nacional" stateProvince="Veracruz">Puente Nacional</location>
,
<collectingDate value="1985-03-13">13 Mar. 1985</collectingDate>
,
<emphasis italics="true" pageId="14" pageNumber="105">
<collectorName>G. Castillo-Campo</collectorName>
&amp;
<collectorName>Medina</collectorName>
4275
</emphasis>
(XAL).
</materialsCitation>
</paragraph>
<paragraph pageId="14" pageNumber="105">
This specimen represents part of the variation found in
<taxonomicName class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia mexicana" order="Cycadales" pageId="14" pageNumber="105" phylum="Tracheophyta" rank="species" species="mexicana">
<emphasis italics="true" pageId="14" pageNumber="105">Ceratozamia mexicana</emphasis>
</taxonomicName>
and the locality is on the collection route that was followed by 19th century collectors to the centre of
<collectingRegion country="Mexico" name="Veracruz de Ignacio de la Llave">Veracruz</collectingRegion>
.
</paragraph>
<paragraph pageId="14" pageNumber="105">
<taxonomicName authorityName="Miq" authorityYear="1847" class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia longifolia" order="Cycadales" pageId="14" pageNumber="105" phylum="Tracheophyta" rank="species" species="longifolia">
<emphasis italics="true" pageId="14" pageNumber="105">Ceratozamia longifolia</emphasis>
</taxonomicName>
.
<materialsCitation collectingDate="1913-04" collectorName="Miq. Tijdschr. Wis-Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch." country="MEXICO" location="Zacuapam" specimenCount="1" stateProvince="Veracruz" typeStatus="Neotype">
<collectorName>Miq. Tijdschr. Wis-Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch.</collectorName>
1 (4): 40. 1848.
<typeStatus>Neotype</typeStatus>
(designated here).
<collectingCountry name="Mexico">MEXICO</collectingCountry>
.
<collectingRegion country="Mexico" name="Veracruz de Ignacio de la Llave">Veracruz</collectingRegion>
:
<location LSID="urn:lsid:plazi:treatment:68E6B809971D5A51BD203414DBAC2F83:51B390CB95C9435D7A7067237FA6CEF8" country="MEXICO" name="Zacuapam" stateProvince="Veracruz">Zacuapam</location>
,
<collectingDate value="1913-04">Apr. 1913</collectingDate>
,
<emphasis italics="true" pageId="14" pageNumber="105">Purpus s/n</emphasis>
(MO [6362]).
</materialsCitation>
</paragraph>
<paragraph pageId="14" pageNumber="105">
This locality is a historic collection from the central region of
<collectingRegion country="Mexico" name="Veracruz de Ignacio de la Llave">Veracruz</collectingRegion>
for
<taxonomicName class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia mexicana" order="Cycadales" pageId="14" pageNumber="105" phylum="Tracheophyta" rank="species" species="mexicana">
<emphasis italics="true" pageId="14" pageNumber="105">Ceratozamia mexicana</emphasis>
</taxonomicName>
, which was one of the most collected cycads during the 19th century.
</paragraph>
<paragraph pageId="14" pageNumber="105">
<taxonomicName authority="Brongn. var. mexicana" class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia mexicana subsp. var. var. mexicana" order="Cycadales" pageId="14" pageNumber="105" phylum="Tracheophyta" rank="variety" species="mexicana" subSpecies="var." variety="mexicana">Ceratozamia mexicana Brongn. var. mexicana</taxonomicName>
.
</paragraph>
<paragraph pageId="14" pageNumber="105">
<taxonomicName authorityName="Miq" authorityYear="1847" class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia longifolia" order="Cycadales" pageId="14" pageNumber="105" phylum="Tracheophyta" rank="species" species="longifolia">
<emphasis italics="true" pageId="14" pageNumber="105">Ceratozamia longifolia</emphasis>
</taxonomicName>
<materialsCitation collectingDate="1975-12-23" collectorName="Miq. Tijdschr. Wis-Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch., M. G. Zola" country="MEXICO" location="Teocelo" specimenCount="1" stateProvince="Veracruz" typeStatus="Neotype">
var.
<taxonomicName class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia longifolia var. minor" order="Cycadales" pageId="14" pageNumber="105" phylum="Tracheophyta" rank="variety" species="longifolia" variety="minor">Ceratozamia longifolia minor</taxonomicName>
.
<collectorName>Miq. Tijdschr. Wis-Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch.</collectorName>
2 (4): 290. 1849.
<typeStatus>Neotype</typeStatus>
(designated here).
<collectingCountry name="Mexico">MEXICO</collectingCountry>
.
<collectingRegion country="Mexico" name="Veracruz de Ignacio de la Llave">Veracruz</collectingRegion>
:
<location LSID="urn:lsid:plazi:treatment:68E6B809971D5A51BD203414DBAC2F83:3E6B84624579A29DC72A06165F5B6338" country="MEXICO" name="Teocelo" stateProvince="Veracruz">Teocelo</location>
,
<collectingDate value="1975-12-23">23 Dic. 1975</collectingDate>
,
<emphasis italics="true" pageId="14" pageNumber="105">
<collectorName>M. G. Zola</collectorName>
146
</emphasis>
(XAL).
</materialsCitation>
</paragraph>
<paragraph pageId="14" pageNumber="105">
This locality is on the historic collection route for
<taxonomicName authorityName="Brongniart" authorityYear="1846" class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia" order="Cycadales" pageId="14" pageNumber="105" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="14" pageNumber="105">Ceratozamia</emphasis>
</taxonomicName>
and represents the northern end of the distribution for
<taxonomicName class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia mexicana" order="Cycadales" pageId="14" pageNumber="105" phylum="Tracheophyta" rank="species" species="mexicana">
<emphasis italics="true" pageId="14" pageNumber="105">Ceratozamia mexicana</emphasis>
</taxonomicName>
where plants with lower leaflet widths have been recorded.
</paragraph>
<paragraph pageId="14" pageNumber="105">
<taxonomicName class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia mexicana subsp. var. var. longifolia" order="Cycadales" pageId="14" pageNumber="105" phylum="Tracheophyta" rank="variety" species="mexicana" subSpecies="var." variety="longifolia">Ceratozamia mexicana var. longifolia</taxonomicName>
. (Miquel) Dyer. Biol. cent.- amer., Bot. 3 (16): 193. 1884.
<typeStatus>Type</typeStatus>
: Based on
<taxonomicName authorityName="Miq" authorityYear="1847" class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia longifolia" order="Cycadales" pageId="14" pageNumber="105" phylum="Tracheophyta" rank="species" species="longifolia">
<emphasis italics="true" pageId="14" pageNumber="105">Ceratozamia longifolia</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection pageId="15" pageNumber="106" type="description">
<paragraph pageId="15" pageNumber="106">
<pageBreakToken pageId="15" pageNumber="106" start="start">Description</pageBreakToken>
.
</paragraph>
<paragraph pageId="15" pageNumber="106">
<emphasis italics="true" pageId="15" pageNumber="106">Stem</emphasis>
epigeous, erect and decumbent, 20-80 cm in length, 20-45 cm in diameter.
<emphasis italics="true" pageId="15" pageNumber="106">Cataphylls</emphasis>
persistent, densely tomentose at emergence, reddish-brown and partially tomentose at maturity, triangular, apex acuminate, 2-6
<normalizedToken originalValue="×">x</normalizedToken>
2.5-6 cm at base.
<emphasis italics="true" pageId="15" pageNumber="106">Leaves</emphasis>
5-55, descending, 100-270 cm, light green at emergence, brown pubescence, glabrous at maturity.
<emphasis italics="true" pageId="15" pageNumber="106">Petiole</emphasis>
terete, straight, 30-93 cm, armed with short and thin prickles, dark green in adult leaves.
<emphasis italics="true" pageId="15" pageNumber="106">Rachis</emphasis>
terete, straight, 56-154 cm, armed with prickles and unarmed, green in adult leaves.
<emphasis italics="true" pageId="15" pageNumber="106">Leaflets</emphasis>
25-42, lanceolate, mostly planar, basally falcate, coriaceous, flat, opposite to subopposite, plane, green, adaxial and abaxial side glabrous, acuminate and symmetric apex, attenuate at base, with conspicuous and light green veins; median leaflets 29-51
<normalizedToken originalValue="×">x</normalizedToken>
2.3-3.7 cm, 1.8-4 cm between leaflets; articulations green, 0.6-1.5 cm wide.
<emphasis italics="true" pageId="15" pageNumber="106">Polliniferous strobilus</emphasis>
generally solitary (1-2), cylindrical, erect, 24-40 cm in length, 5.5-7.6 cm in diameter, greenish-yellow at emergence, greenish-yellow with blackish pubescence at maturity; peduncle tomentose, reddish-brown to light brown, 3.5-5 cm in length, 1.8-2.3 cm in diameter; microsporophylls 1.6-2.7
<normalizedToken originalValue="×">x</normalizedToken>
0.9-2.1 cm, non-recurved distal face.
<emphasis italics="true" pageId="15" pageNumber="106">Ovuliferous strobilus</emphasis>
generally solitary (1-2), cylindrical, erect or pendular, 23.5-38 cm in length, 10.5-14.6 cm in diameter, dark green with blackish pubescence at emergence, dark green with black and grey trichomes at maturity, acuminate apex; peduncle tomentose, brown to reddish-brown, 8-11.5 cm in length, 2.5-3 cm in diameter; megasporophylls 55-224, 2-4
<normalizedToken originalValue="×">x</normalizedToken>
4.4-5.3 cm, prominent distal face, obtuse angle between horns.
<emphasis italics="true" pageId="15" pageNumber="106">Seeds</emphasis>
ovoid, sarcotesta whitish-yellow to yellow when immature, light brown at maturity, 2-3.3 cm in length, 1.5-2.5 cm in diameter.
</paragraph>
</subSubSection>
<subSubSection pageId="15" pageNumber="106" type="distribution">
<paragraph pageId="15" pageNumber="106">Distribution and habitat.</paragraph>
<paragraph pageId="15" pageNumber="106">
Endemic to Mexico from the River La Antigua drainage system within Xico, Teocelo and Coatepec municipalities to south end of Sierra Madre Oriental in Veracruz state, between 500-1,300 m in cloud forest (Figure
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Distribution map of Ceratozamia tenuis, C. totonacorum, C. morettii, C. mexicana, C. delucana, C. decumbens and C. brevifrons." figureDoi="10.3897/phytokeys.100.23152.figure8" httpUri="https://binary.pensoft.net/fig/211379" pageId="15" pageNumber="106">8</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="15" pageNumber="106" type="etymology">
<paragraph pageId="15" pageNumber="106">Etymology.</paragraph>
<paragraph pageId="15" pageNumber="106">The epithet is derived from the country of origin of the material for the description of the genus.</paragraph>
</subSubSection>
<subSubSection pageId="15" pageNumber="106" type="distinguishing features">
<paragraph pageId="15" pageNumber="106">Distinguishing features.</paragraph>
<paragraph pageId="15" pageNumber="106">
<taxonomicName class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia mexicana" order="Cycadales" pageId="15" pageNumber="106" phylum="Tracheophyta" rank="species" species="mexicana">
<emphasis italics="true" pageId="15" pageNumber="106">Ceratozamia mexicana</emphasis>
</taxonomicName>
is distinguished by its lanceolate, coriaceous and flat leaflets with a symmetric apex, ovulate strobilus dark green with black and grey trichomes at maturity, and a prominent distal face with an obtuse angle between horns (Fig.
<figureCitation captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Ceratozamia mexicana. A Adult plant in habit B Leaf at emergence C Detail of pollen strobili D Microsporophyll E Megasporophyll F Detail of ovulate strobili G Seeds." figureDoi="10.3897/phytokeys.100.23152.figure11" httpUri="https://binary.pensoft.net/fig/211382" pageId="15" pageNumber="106">11</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="15" pageNumber="106" type="materials_examined">
<paragraph pageId="15" pageNumber="106">Specimens examined.</paragraph>
<paragraph pageId="15" pageNumber="106">
<collectingCountry name="Mexico">MEXICO</collectingCountry>
.
<emphasis bold="true" pageId="15" pageNumber="106">
<collectingRegion country="Mexico" name="Veracruz de Ignacio de la Llave">Veracruz</collectingRegion>
</emphasis>
: Coatepec,
<emphasis italics="true" pageId="15" pageNumber="106">P. Zamora C. 2450</emphasis>
(MEXU, XAL); Comapa,
<emphasis italics="true" pageId="15" pageNumber="106">
F.
<normalizedToken originalValue="Nicolalde-Morejón">Nicolalde-Morejon</normalizedToken>
et al. 2146-2156
</emphasis>
(CIB),
<emphasis italics="true" pageId="15" pageNumber="106">
L.
<normalizedToken originalValue="Martínez-Domínguez">Martinez-Dominguez</normalizedToken>
et al. 164, 716-730
</emphasis>
(CIB); Puente Nacional,
<emphasis italics="true" pageId="15" pageNumber="106">G. Castillo-Campos &amp; M. E. Medina 4299</emphasis>
(XAL); Sochiapa,
<emphasis italics="true" pageId="15" pageNumber="106">
M.
<normalizedToken originalValue="Vázquez-Torres">Vazquez-Torres</normalizedToken>
8589
</emphasis>
(CIB); Teocelo,
<emphasis italics="true" pageId="15" pageNumber="106">
F.
<normalizedToken originalValue="Nicolalde-Morejón">Nicolalde-Morejon</normalizedToken>
et al. 2273-2278
</emphasis>
(CIB),
<emphasis italics="true" pageId="15" pageNumber="106">M. G. Zola 146</emphasis>
(XAL),
<emphasis italics="true" pageId="15" pageNumber="106">
M.
<normalizedToken originalValue="Vázquez-Torres">Vazquez-Torres</normalizedToken>
</emphasis>
4865 (CIB),
<emphasis italics="true" pageId="15" pageNumber="106">
L.
<normalizedToken originalValue="Martínez-Domínguez">Martinez-Dominguez</normalizedToken>
et al. 731-750, 764-770
</emphasis>
(CIB); Tlaltetela,
<emphasis italics="true" pageId="15" pageNumber="106">
F.
<normalizedToken originalValue="Nicolalde-Morejón">Nicolalde-Morejon</normalizedToken>
&amp; L.
<normalizedToken originalValue="Martínez-Domínguez">Martinez-Dominguez</normalizedToken>
2246, 2253-2256
</emphasis>
(CIB),
<emphasis italics="true" pageId="15" pageNumber="106">
L.
<normalizedToken originalValue="Martínez-Domínguez">Martinez-Dominguez</normalizedToken>
&amp; F.
<normalizedToken originalValue="Nicolalde-Morejón">Nicolalde-Morejon</normalizedToken>
584-586, 620, 628-635
</emphasis>
(CIB); Totutla,
<emphasis italics="true" pageId="15" pageNumber="106">A. P. Vovides 730-733, 748</emphasis>
(XAL),
<emphasis italics="true" pageId="15" pageNumber="106">Brigada T. Walters s/n</emphasis>
(XAL),
<emphasis italics="true" pageId="15" pageNumber="106">
F.
<normalizedToken originalValue="Nicolalde-Morejón">Nicolalde-Morejon</normalizedToken>
et al. 2279-2281
</emphasis>
(CIB),
<emphasis italics="true" pageId="15" pageNumber="106">
F.
<normalizedToken originalValue="Vázquez">Vazquez</normalizedToken>
B. 730
</emphasis>
(XAL),
<emphasis italics="true" pageId="15" pageNumber="106">J. Rees 6344</emphasis>
(IEB),
<emphasis italics="true" pageId="15" pageNumber="106">J. Rees &amp; A. P. Vovides 1660, 1672, 1689</emphasis>
(XAL),
<emphasis italics="true" pageId="15" pageNumber="106">
L.
<normalizedToken originalValue="Martínez-Domínguez">Martinez-Dominguez</normalizedToken>
et al. 704-707, L.
<normalizedToken originalValue="Martínez-Domínguez">Martinez-Dominguez</normalizedToken>
et al. 752-758
</emphasis>
(CIB); Xico,
<emphasis italics="true" pageId="15" pageNumber="106">
L.
<normalizedToken originalValue="Martínez-Domínguez">Martinez-Dominguez</normalizedToken>
&amp; F.
<normalizedToken originalValue="Nicolalde-Morejón">Nicolalde-Morejon</normalizedToken>
640-648
</emphasis>
(CIB).
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="107" type="taxonomic comments">
<paragraph pageId="16" pageNumber="107">
<pageBreakToken pageId="16" pageNumber="107" start="start">Taxonomic</pageBreakToken>
comments.
</paragraph>
<paragraph pageId="16" pageNumber="107">
The taxonomy of
<taxonomicName class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia mexicana" order="Cycadales" pageId="16" pageNumber="107" phylum="Tracheophyta" rank="species" species="mexicana">
<emphasis italics="true" pageId="16" pageNumber="107">Ceratozamia mexicana</emphasis>
</taxonomicName>
has undergone constant changes since its original publication in 1846. The identity of this binomial was ambiguous for several decades. This taxonomic uncertainty was due to the complexity of relationships with some of the other names published by Miquel during the immediately succeeding years (1847, 1848 and 1849), following the original publication of the genus -namely,
<taxonomicName lsidName="C. brevifrons" pageId="16" pageNumber="107" rank="species" species="brevifrons">
<emphasis italics="true" pageId="16" pageNumber="107">C. brevifrons</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="C. robusta" pageId="16" pageNumber="107" rank="species" species="robusta">
<emphasis italics="true" pageId="16" pageNumber="107">C. robusta</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="C. intermedia" pageId="16" pageNumber="107" rank="species" species="intermedia">
<emphasis italics="true" pageId="16" pageNumber="107">C. intermedia</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="C. longifolia" pageId="16" pageNumber="107" rank="species" species="longifolia">
<emphasis italics="true" pageId="16" pageNumber="107">C. longifolia</emphasis>
</taxonomicName>
, and
<taxonomicName lsidName="C. latifolia" pageId="16" pageNumber="107" rank="species" species="latifolia">
<emphasis italics="true" pageId="16" pageNumber="107">C. latifolia</emphasis>
</taxonomicName>
. Later on, some names were placed in synonymy and others were characterised as
<emphasis italics="true" pageId="16" pageNumber="107">nomina nuda</emphasis>
; in particular,
<taxonomicName lsidName="C. brevifrons" pageId="16" pageNumber="107" rank="species" species="brevifrons">
<emphasis italics="true" pageId="16" pageNumber="107">C. brevifrons</emphasis>
</taxonomicName>
was listed as a synonym of
<taxonomicName lsidName="C. mexicana" pageId="16" pageNumber="107" rank="species" species="mexicana">
<emphasis italics="true" pageId="16" pageNumber="107">C. mexicana</emphasis>
</taxonomicName>
under the assumption that it was in fact a juvenile form of the latter (see
<bibRefCitation author="Miquel, FAW" journalOrPublisher="Tijschrift voor de Wis - en Natuurkundige Wetenschappen" pageId="25" pageNumber="116" pagination="197 - 208" refId="B19" refString="Miquel, FAW, 1848. Over eenige nieuwe of zeldame Cycadeen in den Hortus Botanicus te Amsterdam. Derde gedeelte. Tijschrift voor de Wis -en Natuurkundige Wetenschappen 1 (4): 197 - 208" title="Over eenige nieuwe of zeldame Cycadeen in den Hortus Botanicus te Amsterdam. Derde gedeelte." volume="1" year="1848">Miquel 1848</bibRefCitation>
,
<bibRefCitation author="Miquel, FAW" journalOrPublisher="van der Post, 1861" pageId="26" pageNumber="117" refId="B21" refString="Miquel, FAW, 1861. Prodromus systematis Cycadearum. In honorem festi diei XV kal. m. Julii MDCCCLXI, quo academia rheno-trajectina exacta XLV lustra celebrat. Utrecht, C. van der. Post Jr., Amsterdam, C. G. van der Post, 1861" title="Prodromus systematis Cycadearum. In honorem festi diei XV kal. m. Julii MDCCCLXI, quo academia rheno-trajectina exacta XLV lustra celebrat. Utrecht, C. van der. Post Jr., Amsterdam, C. G." year="1861">1861</bibRefCitation>
).
</paragraph>
<paragraph pageId="16" pageNumber="107">
In further taxonomic treatments of
<taxonomicName authorityName="Brongniart" authorityYear="1846" class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia" order="Cycadales" pageId="16" pageNumber="107" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="16" pageNumber="107">Ceratozamia</emphasis>
</taxonomicName>
-specifically, in
<bibRefCitation author="De Candolle, A" journalOrPublisher=")" pageId="25" pageNumber="116" refId="B8" refString="De Candolle, A, 1864-1868. Prodromus Systematis Universalis Regni Vegetabilis Pars XVI Sectio Posterior, 546-547." title="Prodromus Systematis Universalis Regni Vegetabilis Pars XVI Sectio Posterior, 546 - 547." year="1864 - 1868">De Candolle (1864-1868)</bibRefCitation>
and including
<normalizedToken originalValue="Miquels">Miquel's</normalizedToken>
<emphasis italics="true" pageId="16" pageNumber="107">
Nouveaux
<normalizedToken originalValue="matériaux">materiaux</normalizedToken>
pour servir
<normalizedToken originalValue="à">a</normalizedToken>
la connaissance des
<normalizedToken originalValue="Cycadées">Cycadees</normalizedToken>
</emphasis>
(1868-1869a, b)- this nomenclatural status was maintained. In his mature work, Miquel considered the variation observed between juveniles and adults was due to the corresponding ontogenetic modifications and collapsed into synonymy with
<taxonomicName lsidName="C. mexicana" pageId="16" pageNumber="107" rank="species" species="mexicana">
<emphasis italics="true" pageId="16" pageNumber="107">C. mexicana</emphasis>
</taxonomicName>
all five species previously described.
</paragraph>
<paragraph pageId="16" pageNumber="107">
Half a century later,
<bibRefCitation author="Schuster, J" journalOrPublisher="Acta Botanica Mexicana" pageId="27" pageNumber="118" refId="B40" refString="Schuster, J, 1932. Cycadaceae. In: Engler A (Ed.) Das Pflanzenreich 99: 1-168." title="Cycadaceae. In: Engler A (Ed.) Das Pflanzenreich 99: 1 - 168." year="1932">Schuster (1932)</bibRefCitation>
circumscribed the genus to include only two species:
<taxonomicName class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia kuesteriana" order="Cycadales" pageId="16" pageNumber="107" phylum="Tracheophyta" rank="species" species="kuesteriana">
<emphasis italics="true" pageId="16" pageNumber="107">Ceratozamia kuesteriana</emphasis>
</taxonomicName>
and
<taxonomicName lsidName="C. mexicana" pageId="16" pageNumber="107" rank="species" species="mexicana">
<emphasis italics="true" pageId="16" pageNumber="107">C. mexicana</emphasis>
</taxonomicName>
, with two varieties and two forms. However, in the first modern taxonomic treatment (
<bibRefCitation author="Vovides, A" journalOrPublisher="Botany" pageId="27" pageNumber="118" refId="B45" refString="Vovides, A, Rees, JD, Vazque-Torres, M, 1983. Zamiaceae. Fasciculo 26. Flora de Veracruz. Xalapa, Veracruz, 1-11." title="Zamiaceae. Fasciculo 26. Flora de Veracruz. Xalapa, Veracruz, 1 - 11." year="1983">Vovides et al. 1983</bibRefCitation>
) and continuing with the typification of names within the genus (
<bibRefCitation DOI="https://doi.org/10.2307/1221921" author="Stevenson, DW" journalOrPublisher="Taxon" pageId="27" pageNumber="118" pagination="578 - 584" refId="B42" refString="Stevenson, DW, Sabato, S, 1986. Typification of names in Ceratozamia Brongn. Dion Lindl., and Microcycas A. D.C. (Zamiaceae). Taxon 35 (3): 578 - 584, DOI: https://doi.org/10.2307/1221921" title="Typification of names in Ceratozamia Brongn. Dion Lindl., and Microcycas A. D. C. (Zamiaceae)." url="https://doi.org/10.2307/1221921" volume="35" year="1986">Stevenson and Sabato 1986</bibRefCitation>
), the type species was circumscribed to the central region of Veracruz. In this context,
<bibRefCitation author="Vovides, A" journalOrPublisher="Botany" pageId="27" pageNumber="118" refId="B45" refString="Vovides, A, Rees, JD, Vazque-Torres, M, 1983. Zamiaceae. Fasciculo 26. Flora de Veracruz. Xalapa, Veracruz, 1-11." title="Zamiaceae. Fasciculo 26. Flora de Veracruz. Xalapa, Veracruz, 1 - 11." year="1983">Vovides et al. (1983)</bibRefCitation>
attributed a wide geographic distribution and high variation of morphological characters to
<taxonomicName lsidName="C. mexicana" pageId="16" pageNumber="107" rank="species" species="mexicana">
<emphasis italics="true" pageId="16" pageNumber="107">C. mexicana</emphasis>
</taxonomicName>
while recognising three varieties:
<taxonomicName authority="var. mexicana" lsidName="C. mexicana subsp. var. var. mexicana" pageId="16" pageNumber="107" rank="variety" species="mexicana" subSpecies="var." variety="mexicana">C. mexicana var. mexicana</taxonomicName>
,
<taxonomicName lsidName="C. mexicana subsp. var. var. latifolia" pageId="16" pageNumber="107" rank="variety" species="mexicana" subSpecies="var." variety="latifolia">C. mexicana var. latifolia</taxonomicName>
(Miq.) Schuster and
<taxonomicName lsidName="C. mexicana subsp. var. var. robusta" pageId="16" pageNumber="107" rank="variety" species="mexicana" subSpecies="var." variety="robusta">C. mexicana var. robusta</taxonomicName>
(Miq.) Dyer.
</paragraph>
<paragraph pageId="16" pageNumber="107">
The original
<taxonomicName class="Cycadopsida" family="Zamiaceae" genus="Ceratozamia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ceratozamia mexicana" order="Cycadales" pageId="16" pageNumber="107" phylum="Tracheophyta" rank="species" species="mexicana">
<emphasis italics="true" pageId="16" pageNumber="107">Ceratozamia mexicana</emphasis>
</taxonomicName>
specimens were collected by the Belgian botanist and explorer A. B. Ghiesbreght, who probably did his fieldwork in the region of Huatusco, Veracruz. This location is recognised as part of his route for botanical collections between 1835 and 1838 (
<bibRefCitation author="Barnhart, JH" pageId="24" pageNumber="115" refId="B1" refString="Barnhart, JH, 1965. Biographical Notes upon Botanists. In: Hall GK (Ed.) New York Botanical Garden. Boston." title="Biographical Notes upon Botanists. In: Hall GK (Ed.) New York Botanical Garden. Boston." year="1965">Barnhart 1965</bibRefCitation>
;
<bibRefCitation author="Sartorius, CC" journalOrPublisher="Acta Botanica Mexicana" pageId="26" pageNumber="117" refId="B39" refString="Sartorius, CC, 1990. Mexico hacia 1850. Estudio preliminar, revision y notas Brigida Von Mentz. Consejo Nacional para la Cultura y las Artes. Mexico, D.F., 1-327." title="Mexico hacia 1850. Estudio preliminar, revision y notas Brigida Von Mentz. Consejo Nacional para la Cultura y las Artes. Mexico, D. F., 1 - 327." year="1990">Sartorius 1990</bibRefCitation>
;
<bibRefCitation author="Rzedowski, J" journalOrPublisher="Acta Botanica Mexicana" pageId="26" pageNumber="117" refId="B38" refString="Rzedowski, J, Calderon, G, Butanda, A, 2009. Los principales colectores de plantas activos en Mexico entre 1700 y 1930. Instituto de Ecologia, A. C. y Comision Nacional para el Conocimiento y Uso de la Biodiversidad. Patzcuaro, Michoacan, 1-144." title="Los principales colectores de plantas activos en Mexico entre 1700 y 1930. Instituto de Ecologia, A. C. y Comision Nacional para el Conocimiento y Uso de la Biodiversidad. Patzcuaro, Michoacan, 1 - 144." year="2009">Rzedowski et al. 2009</bibRefCitation>
). Besides the &quot;El Mirador&quot; locality, mentioned by
<bibRefCitation author="Thiselton-Dyer, WTT" journalOrPublisher="Botany" pageId="27" pageNumber="118" pagination="190 - 195" refId="B44" refString="Thiselton-Dyer, WTT, 1884. Order CXXXVII. Cycadaceae. Biologia Centrali-Americana. Botany 3: 190 - 195" title="Order CXXXVII. Cycadaceae. Biologia Centrali-Americana." volume="3" year="1884">Thiselton-Dyer (1884)</bibRefCitation>
and
<bibRefCitation author="Schuster, J" journalOrPublisher="Acta Botanica Mexicana" pageId="27" pageNumber="118" refId="B40" refString="Schuster, J, 1932. Cycadaceae. In: Engler A (Ed.) Das Pflanzenreich 99: 1-168." title="Cycadaceae. In: Engler A (Ed.) Das Pflanzenreich 99: 1 - 168." year="1932">Schuster (1932)</bibRefCitation>
as the main reference place, the species still exists in Huatusco. Due to coffee cultivation, the original vegetation has been replaced almost in its entirety. However, the species still occurs in the wild in relictual secondary vegetation within a property called &quot;Hacienda Zacuapam&quot;. This property was part of a former &quot;El Mirador&quot; hacienda, early in the XIX century (
<bibRefCitation author="Sartorius, CC" journalOrPublisher="Acta Botanica Mexicana" pageId="26" pageNumber="117" refId="B39" refString="Sartorius, CC, 1990. Mexico hacia 1850. Estudio preliminar, revision y notas Brigida Von Mentz. Consejo Nacional para la Cultura y las Artes. Mexico, D.F., 1-327." title="Mexico hacia 1850. Estudio preliminar, revision y notas Brigida Von Mentz. Consejo Nacional para la Cultura y las Artes. Mexico, D. F., 1 - 327." year="1990">Sartorius 1990</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>