Evolutionary Genetics of the Genus Zamia (Zamiaceae, Cycadales)

The genus Zamia L. (Zamiaceae), consisting of 77 species, is the most species-rich and widely distributed cycad genus in the New World and is arguably the most morphologically and ecologically diverse genus in the Cycadales. We utilized a multilocus sequence dataset of 10 independent loci (9 single copy nuclear genes + 1 plastid) and extensive taxon sampling (over 90% of species) to infer phylogenetic relationships within Zamia. We infer a robust phylogenetic tree for the genus with a strong geographic delimitation of clades and find that four morphological characters typically used for diagnostic purposes in the genus exhibit a high degree of homoplasy. We genotyped four populations of the Belizean endemic cycad Zamia decumbens using ten microsatellite loci and analyzed the data using a variety of population genetic analyses methods. The populations occurred in two different habitat types: inside dolines (one at a cave entrance and two at the bottom of sinkholes) and one on steep karstic terrain on a hilltop. We found the genetic variation was not structured geographically or by habitat type, but rather seemed to reflect the demographic history of the populations and their genetic connectivity. Contemporary geneflow between populations is generally low, with the Cave population being the most important population in facilitating genetic con­nectivity in the region, mostly as a source of migrants to other populations. A conservation assessment for the three cycad species native to the Bahamas Islands is presented based on field surveys on all islands where these species occur. Zamia angustifolia is native to Eleuthera, Zamia integrifolia is native to Abaco, Andros, Eleuthera, Grand Bahama and New Providence, and Zamia lucayana is endemic to Long Island. We assessed the genetic structure of Z. lucayana based on 15 polymorphic microsatellite DNA loci; this indicated that the three known populations should be considered a single management unit. However, the high number of private alleles suggests that genetic drift, indicative of recent fragmentation, is progressing. We propose in situ conservation strategies, and we also collected germplasm from a total of 24 populations of these three cycad species, for ex situ conservation

Propagation of Cycad Collections from Seed

Propagation of cycads from seed can aid their conservation as it helps reduce the demand for wildcollected plants. Seed-produced plants can be used for reintroduction programmes if the parent plants are from known provenance and care is taken to avoid hybridisation. This paper discusses the techniques required for successful seed propagation of cycads, including pollen collection, storage, viability testing, manual pollination, seed collection, storage and germination

Molecular and Morphological Phylogenetic Analyses of New World Cycad Beetles: What They Reveal about Cycad Evolution in the New World

Two major lineages of beetles inhabit cycad cones in the New World: weevils (Curculionoidea) in the subtribe Allocorynina, including the genera Notorhopalotria Tang and O’Brien, Parallocorynus Voss, Protocorynus O’Brien and Tang and Rhopalotria Chevrolat, and beetles in the family Erotylidae, including the genus Pharaxonotha Reitter. Analysis of the 16S ribosomal RNA (rRNA) mitochondrial gene as well as cladistic analysis of morphological characters of the weevils indicate four major radiations, with a probable origin on the cycad genus Dioon Lindl. and comparatively recent host shifts onto Zamia L. Analysis of the 16S rRNA gene for erotylid beetles indicates that an undescribed genus restricted to New World Ceratozamia Brongn. is the most early-diverging clade, and this lineage is sister to a large radiation of the genus Pharaxonotha onto Zamia, with apparent host shifts onto Dioon and Ceratozamia. Analysis of beetles are in accord with current models of continental drift in the Caribbean basin, support some proposed species groupings of cycads, but not others, and suggest that pollinator type may impact population genetic structure in their host cycads

A NEW CLIFF-DWELLING SPECIES OF ZAMIA (ZAMIACEAE) FROM BELIZE

Volume: 3Start Page: 23End Page: 2

Evolutionary Genetics of the Genus Zamia (Zamiaceae, Cycadales)

The genus Zamia L. (Zamiaceae), consisting of 77 species, is the most species-rich and widely distributed cycad genus in the New World and is arguably the most morphologically and ecologically diverse genus in the Cycadales. We utilized a multilocus sequence dataset of 10 independent loci (9 single copy nuclear genes + 1 plastid) and extensive taxon sampling (over 90% of species) to infer phylogenetic relationships within Zamia. We infer a robust phylogenetic tree for the genus with a strong geographic delimitation of clades and find that four morphological characters typically used for diagnostic purposes in the genus exhibit a high degree of homoplasy. We genotyped four populations of the Belizean endemic cycad Zamia decumbens using ten microsatellite loci and analyzed the data using a variety of population genetic analyses methods. The populations occurred in two different habitat types: inside dolines (one at a cave entrance and two at the bottom of sinkholes) and one on steep karstic terrain on a hilltop. We found the genetic variation was not structured geographically or by habitat type, but rather seemed to reflect the demographic history of the populations and their genetic connectivity. Contemporary geneflow between populations is generally low, with the Cave population being the most important population in facilitating genetic con­nectivity in the region, mostly as a source of migrants to other populations. A conservation assessment for the three cycad species native to the Bahamas Islands is presented based on field surveys on all islands where these species occur. Zamia angustifolia is native to Eleuthera, Zamia integrifolia is native to Abaco, Andros, Eleuthera, Grand Bahama and New Providence, and Zamia lucayana is endemic to Long Island. We assessed the genetic structure of Z. lucayana based on 15 polymorphic microsatellite DNA loci; this indicated that the three known populations should be considered a single management unit. However, the high number of private alleles suggests that genetic drift, indicative of recent fragmentation, is progressing. We propose in situ conservation strategies, and we also collected germplasm from a total of 24 populations of these three cycad species, for ex situ conservation

Stem Branching of Cycad Plants Informs Horticulture and Conservation Decisions

The number of branches in male and female plants of Cycas micronesica K.D. Hill, Cycas edentata de Laub., Cycas wadei Merr., and Zamia encephalartoides D.W. Stev. were counted in Guam, Philippines, and Colombia, to confirm earlier reports that female plants develop fewer branches than males. Cycas plants produce determinate male strobili and indeterminate female strobili, but Zamia plants produce determinate strobili for both sexes. More than 80% of the female trees for each of the Cycas species were unbranched with a single stem, but more than 80% of the male trees exhibited two or more branches. The mean number of branches on male plants was more than double that of female plants. The number of branches of the Zamia male plants was almost triple that of female plants. Moreover, the Zamia plants produced 2.8-fold greater numbers of branches than the mean of the Cycas plants. Most of Guam’s unsexed C. micronesica trees in 2004 were unbranched, but after 15 years of damage from non-native insect herbivores, most of the remaining live trees in 2020 contained three or more branches. The results confirm that male Cycas and Zamia plants produce more branches than female plants and suggest cycad species with determinate female strobili produce more branches on female plants than species with indeterminate female strobili. Our results indicate that the years of plant mortality on Guam due to non-native insect herbivores have selectively killed more female C. micronesica trees. Horticulture and conservation decisions may be improved with this sexual dimorphism knowledge

WHAT IS ZAMIA PRASINA (ZAMIACEAE: CYCADALES)?

Volume: 3Start Page: 43End Page: 4

A review of the Cenozoic fossil record of the genus Zamia L. (Zamiaceae, Cycadales) with recognition of a new species from the late Eocene of Panama – evolution and biogeographic inferences

Modern Zamia L. is the second largest genus among cycads, however reliably identified fossil occurrences of the genus have so far been missing. Previously, fossil “Zamia” species were established in large numbers on the basis of macromorphological similarity of foliage fragments to living Zamia species. However, a reinvestigation of specimens assigned formerly to Zamia and the relevant literature provided no clear-cut evidence for their assignment to this genus. We investigated a newly recovered fossil specimen from marine sediments of the Gatuncillo Formation, near Buena Vista, Colon Province, Central Panama. It represents the first unequivocal fossil record of the genus confirmed by epidermal as well as macromorphological characters and it is described as Zamia nelliae Erdei & Calonje sp. nov. Foraminiferal and nannoplankton biostratigraphy of the locality indicates a late Eocene to earliest Oligocene age. Morphometric comparison of epidermal features of Z. nelliae with those of modern Zamia species suggests similarity with those of the Caribbean Zamia clade. The fossil record of Zamia from Panama implies that the genus appeared by the end of the Eocene or earliest Oligocene in the Central American–Caribbean region, however, the origin of the genus is still unresolved. The record of Z. nelliae may challenge former concepts on the evolution of Zamia and raises an “intermediate” hypothesis on its origin in the Central American–Caribbean region and its subsequent dispersal south- and northwards. • Keywords: cycad, Zamia, cuticle, morphometry, Paleogene, Panama

Evolutionary signal of leaflet anatomy in the Zamiaceae

Premise of research: The morphology of leaves is shaped by both historical and current selection acting on constrained developmental systems. For this reason, the phylogenetic signal of these characters is usually overlooked. Methodology: We investigate morphology of the leaflets of all genera of the Zamiaceae using multiple microscopical techniques to test whether leaf characters present a phylogenetic signal and whether they are useful to define clades at a suprageneric level. Pivotal results: Our investigation shows that most genera are quite uniform in their leaflet anatomy, with the largest genera (Zamia, Encephalartos) presenting the highest degree of variation. Using both Bayesian and parsimony methods on two different molecular scaffolds, we are able to show that leaflet anatomy has a strong phylogenetic signal in the Zamiaceae and that many clades retrieved by molecular analyses present potential synapomorphies in their leaflet anatomy. Particularly, the placement of Stangeria in a clade with Zamia and Microcycas is supported by the presence of both an adaxial and an abaxial girder sclerenchyma and the absence of sclerified hypodermis. The placement of Stangeria as sister to Bowenia, on the other hand, is not supported by our analysis. Instead, our results put into question the homology of the similar guard cell morphology in the two genera. Conclusions: We show that leaflet anatomy has a substantial amount of phylogenetic signal in the Zamiaceae, supporting relationships that are not supported by general morphology. Therefore, anatomical investigation represents a promising avenue for plant systematists

A new arborescent species of zamia (cycadales, zamiaceae) from the department of huila, eastern cordillera of colombia

Zamia huilensis Calonje, Esquivel, and amp; D.W. Stev., a new arborescent species from theEastern Cordillera of Colombia (Department of Huila), is described and illustrated. Itis compared to Z. muricata Willd., which has similar leaflet shape, and to Z. tolimensisCalonje, Esquivel and amp; D.W. Stev., with which it shares some reproductive characters.It differs from Z. muricata in having an arborescent trunk, beige-yellow to beigeovulate strobili supported by short peduncles, and microsporangia aggregated intoa single group on abaxial side of microsporophyll; it differs from Z. muricata in thesubterranean trunk, dark brown to black ovulate strobili supported by long peduncles,and microsporangia separated into two groups on abaxial side of microsporophylls.It differs from Z. tolimensis in having an overall smaller size, leaflets that are notstrongly falcate and with margins toothed below the distal half, compared to Z. tolimensiswhich is a much larger plant, and has strongly falcate leaflets that are nottoothed below the distal half