Karyotype evolution by centromeric fission inZamia (Cycadales)

The chromosome numbers of several species ofZamia from Mexico are reported.Z. paucijuga, distributed from central Oaxaca to Nayarit, has been found to have 2n = 23, 25, 26, 27 and 28. 2n = 28 is the highest chromosome number yet found in the cycads. Karyotypes of this species differ principally in the number of telocentric and metacentric chromosomes present in each; 2n = 23, 25, 26, 27 and 28 were found to have 5, 3, 2, 1 and 0 metacentric and 8, 12, 14, 16 and 18 telocentric chromosomes, respectively.Z. fischeri has been found to be 2n = 16,Z. furfuracea andZ. loddigesii 2n = 18.Zamia paucijuga on the basis of morphological and ecological characteristics, is considered to be an advanced member of this genus. Chromosome and karyotype evolution inZ. paucijuga may have occurred by centromeric fission of metacentric chromosomes; the karyotypes ofZ. paucijuga are strongly asymmetrical, suggesting that they evolved recently.     

Organization of a dispersed repeated DNA element in the Zamia genome

Occurrence and genomic organization of dispersed elements containing ZpS1 satellite repeats have been investigated in a wide representation of species of the old plant genus Zamia (Zamiaceae, Cycadales). In Z. paucijuga, the ZpS1 repeat is organized as long satellite DNA arrays and as short arrays inserted into AT-rich dispersed elements. A comparative study by Southern analysis shows that these unusual dispersed elements containing the ZpS1 repeat are present with different organizations in all investigated Zamia species. In some species these elements are present with a low copy number, while in other species secondary amplification events, involving specific sequence clusters, appear to have generated characteristic dispersed elements in a high copy number. Among Zamia species, several groups share similar restriction patterns, as the Zamia loddigesii complex and the Caribbean species suggesting a general correlation between organization and genomic representation of the dispersed repeated sequence and the pattern of phyletic relationships in the genus. However, the finding of different patterns also among closely related species suggests a complex history of amplifications and losses of these dispersed repetitive elements that cannot be always easily traced through the phylogenetic reconstruction of this ancient plant group.     

A new species of <Emphasis Type="Italic">Zamia</Emphasis> from Chocó, Colombia (Cycadales,Zamiaceae)

Zamia pyrophylla, a new species from Chocó, Colombia, is described and illustrated. It is compared to Z. amazonum, Z. cunaria, and Z. ipetiensis. It is distinguished by its bright orange and red emerging leaves with leaflets that progressively turn green from the apex to the base as they mature, petioles with branched prickles and with ferruginous pubescence when immature, villous strobilar axes, and adaxial microsporangia.     

Chromosome phylogeny of <Emphasis Type="Italic">Zamia</Emphasis> and <Emphasis Type="Italic">Ceratozamia</Emphasis> by means of Robertsonian changes detected by fluorescence <Emphasis Type="Italic">in situ</Emphasis> hybridization (FISH) technique of rDNA

 Appearance and location of 45S rDNA and 5S rDNA signals were compared in chromosomes of nine species of the aneuploid Zamia and their taxonomically and phylogenetically closely related Ceratozamia mexicana. The 45S rDNA signal was detected in the proximal region of six chromosomes in Zamia angustifolia, Z. integrifolia, Z. pumila and Z. pygmaea (all 2n=16); in the proximal region of 6–14 chromosomes in Z. furfuracea, Z. loddigesii, Z. skinneri and Z. vazquezii (all 2n=18); and on the proximal region of 20 chromosomes in Z. muricata (2n=23). The 5S rDNA signals were commonly seen near the terminal region of the short arm of two metacentric chromosomes in the four species with 2n=16 and Z. furfuracea, Z. loddigesii and Z. vazquezii with 2n=18. Other 5S rDNA signals were seen near the terminal region of two terminal-centromeric chromosomes in Z. skinneri and near the terminal region of a metacentric and a telocentric chromosomes in Z. muricata. In contrast, those with 45S and 5S rDNA signals were exhibited in chromosomes of Ceratozamia mexicana in a different manner from those in the nine species of Zamia; the 45S rDNA signal in the terminal region of four metacentric and two submetacentric chromosomes and the 5S rDNA signal near the proximal region of two metacentric chromosomes. Received November 1, 1999 Accepted January 10, 2001     

The identity of <Emphasis Type="Italic">Zamia katzeriana</Emphasis> and <Emphasis Type="Italic">Z. verschaffeltii</Emphasis> (Zamiaceae)

The morphological variation of recently described species (also the older taxa) of Zamia distributed within southeastern Mexico that have wide, coriaceous leaflets is analyzed. The complex is designated as the Z. katzeriana species complex in reference to an historic collection of this name, which is also the earliest named species in the complex. The complex consists of Z. cremnophila, Z. lacandona, Z. splendens and Z. purpurea. During the 1990’s, Z. splendens was separately synonymized under Z. katzeriana and Z. verschaffeltii, both collected in Mexico by German collectors during the 18th century, but the precise locality is unknown. Information concerning the incomplete holotype of Z. verschaffeltii is particularly ambiguous, and the possible type locality suggested by Schuster, Socorro, is imprecise, thus generating taxonomic confusion. Morphometric characterization and discriminant analysis of the contemporary and historic collections (i.e., Z. katzeriana and Z. verschaffeltii) included all the known populations (11) and individuals (115) of the complex throughout its range. The results show that Z. verschaffeltii is not morphometrically related to any of the species in the complex and that Z. splendens should be considered a synonym of Z. katzeriana.

---

Resumen  En este artículo se caracteriza la variación morfológica de las especies de un complejo de Zamia, que presentan folíolos anchos y coriáceos con distribución en el sureste de México, la mayoría de reciente descripción. El complejo es denominado Z. katzeriana, por una colección histórica de este nombre, y consiste de Z. cremnophila, Z. lacandona, Z. splendens y Z. purpurea. Zamia splendens fue sinonimizada separadamente bajo dos colecciones históricas Z. katzeriana y Z. verschaffeltii, de las cuales se desconoce con exactitud su lugar de procedencia, excepto que fueron colectadas en México por botánicos alemanes del siglo XVIII. La información, en especial sobre el holotipo incompleto de Z. verschaffeltii es muy ambigua y la posible localidad tipo mencionada por Schuster, Socorro, es imprecisa, lo cual ha generado confusiones taxonómicas. La caracterización de la variación morfométrica incluyó a todas las poblaciones (11) e individuos (115) conocidas actualmente para el complejo en todo su rango de distribución en México. El análisis discriminante incluyendo a las colecciones históricas, determina que Z. verschaffeltii no está relacionado morfométricamente con ninguna especie del complejo y Z. katzeriana está estrechamente correlacionada con Z. splendens.

     

Zapoteca quichoi (Leguminosae, Mimosoideae), a new species from southeastern Mexico

Zapoteca quichoi, a new species endemic to southern Tabasco, Mexico, is described and illustrated. This species exhibits morphological characteristics that place it within subgenus Amazonica, together with Z. amazonica and Z. microcephala, from Peru and Colombia, respectively.

---

Resumen  Se describe e ilustra Zapoteca quichoi (Leguminosae, Mimosoideae), una nueva especie de Tabasco, México. Esta especie exhibe características morfológicas que la ubican dentro del subgénero Amazonica, junto con Z. amazonica y Z. microcephala, especies nativas de Perú y Colombia respectivamente.

     

Zamia grijalvensis sp. nov. (Zamiaceae,Cycadales) from Chiapas,Mexico with notes on hybridization and karyology

Zamia grijalvensis, a new cycad species from Chiapas, México, is described and illustrated. This species is similar to Z. lacandona with respect to leaves, but differs in habit, morphology and colour of the female and male cones. The female cone of Z. grijalvensis is reddish brown to orange brown, whereas that of Z. lacandona is dark brown with lighter beige tomentum. The male cone of Z. grijalvensis is erect, whereas that of Z. lacandona is decumbent. The new species may be of natural hybrid origin and is discussed in the light of other findings among cycads. Preliminary diploid chromosome counts for Z. grijalvensis 2n = 19, 20 and karyotypes appear variable, as previously found in its congeners with wide distribution, such as Z. paucijuga and Z. prasina, both of which are characterised by high diploid numbers with a high number of telocentric and few metacentric chromosomes. These changes in chromosome number are probably the result of stressful environmental factors.     

Genome sizes for 71 species of Zamia (Cycadales: Zamiaceae) correspond with three different biogeographic regions

Nuclear DNA contents (2C‐value) are reported for 71 out of 76 accepted species of Zamia (Zamiaceae) using flow cytometry with propidium iodide. Nuclear DNA content in Zamia ranges between 33.7 and 45.7 pg. Despite this small range, the largest genome contains roughly 10¹⁰ more base pairs than the smallest genome. The results for Zamia point to two centers of biogeographic distribution: Mexico and Colombia. Nicaragua seems to be the biogeographic boundary for these two centres for Zamia. To the north, genome sizes of 33.7–38.0 pg (average 35.6 pg) are found and to the south (Costa Rica, Panama and South America) 41.2–45.7 pg (average 42.9 pg). Plants from the Caribbean islands (including Florida) have intermediate genome sizes with 37.3–40.9 pg (average 38.7 pg). Costa Rica and Panama are in a transition zone and its species can be divided into three subsections: four species with ‘Caribbean’ values of 38.4–39.5 pg (average 39.0 pg), six species with ‘South American’ values with 42.7–43.6 pg, (average 42.9 pg, and six species with intermediate values ranging between 40.1–41.0 pg (average 40.4 pg). The latter values are nearly absent in other areas, suggesting that they could be the products of (introgressive) hybridization. This study represents the first, nearly complete overview of the genome sizes of the genus Zamia and their relationship with biogeography.     

KARYOTYPIC DATA ON NORTH AND CENTRAL AMERICAN ZAMIACEAE (CYCADALES) AND THEIR PHYLOGENETIC IMPLICATIONS

Chromosome numbers and karyotypes of species from four American Zamiaceae (Cycadales) are reported. Zamia shows interspecific and intraspecific chromosome variation, whereas Microcycas, Ceratozamia, and Dioon have constant karyotypes within each genus. In Zamia, all karyotypes have the same number of submetacentric and acrocentric chromosomes, but they differ in the number of metacentric and telocentric chromosomes. Centric fission of metacentric chromosomes is proposed to explain the karyotypic variation in this genus. Zamia shows karyological relationships with Microcycas and Ceratozamia, whereas Dioon appears very distinct from the other American cycad genera. Affinity among Zamia, Ceratozamia, and Microcycas karyotypes and distinctiveness of Dioon karyotypes are supported by comparative analysis of phenotypic characters in the four genera.     

A character‐based approach in the Mexican cycads supports diverse multigene combinations for DNA barcoding

A DNA barcoding study was conducted to determine the optimal combination of loci needed for successful species‐level molecular identification in three extant cycad genera—Ceratozamia, Dioon, and Zamia—that occur in Mexico. Based on conclusions of a previous multigene study in representative species of all genera in the Cycadales, we tested the DNA barcoding performance of seven chloroplast coding (matK, rpoB, rpoC1, and rbcL) and non‐coding (atpF/H, psbK/I, and trnH‐psbA) regions, plus sequences of the nuclear internal transcribed spacer. We analysed data under the assumptions of the “character attributes organization system” (CAOS), a character‐based approach in which species are identified through the presence of ‘DNA diagnostics’. In Ceratozamia, four chloroplast regions and one nuclear region were needed to achieve > 70% unique species identification. In contrast, the two‐gene combination atpF/H + psbK/I and the four‐gene combination atpF/H + psbK/I + rpoC1 + ITS2 were needed to reach 79% and 75% unique species identification in Dioon and Zamia, respectively. The combinations atpF/H + psbK/I and atpF/H + psbK/I + rpoC1 + ITS2 include loci previously considered by the international DNA barcoding community. However, none of the three combinations of potential DNA barcoding loci found to be optimal with a character‐based approach in the Mexican cycads coincides with the ‘core barcode’ of chloroplast markers (matK + rbcL) recently proposed for universal use in the plant kingdom.     

Karyology and phylogeny of some Mesoamerican species of Zamia (Zamiaceae)

Chromosome numbers and karyotypes of four species of Zamia L. (Zamiaceae) are described. Plants of Z. manicata from Colombia are 2n = 18 with eight metacentric (M), four submetacentric (S), two acrocentric (A), and four telocentric (T) chromosomes. Plants of Z. ipetiensis from Panama are 2n = 23 with 3M + 4S + 2A + 14T. Plants of Z. cunaria from Panama have two different chromosome numbers, 2n = 23 with 3M + 4S + 2A + 14T and 2n = 24 with 2M + 4S + 2A + 16T. Plants of Z. acuminata from Costa Rica and Panama are 2n = 24 with 2M + 4S + 2A + 16T. On the basis of the occurrence of a one-to-two-ratio in the variation of M- and T-chromosome numbers in the karyotypes, centric fission or fusion are considered for their potential involvement in the chromosome variation of these plants. Data deriving from morphology and karyology, interpreted in a cladistic framework, suggest that centric fission rather than centric fusion is involved in the karyotype diversification of the four species and their closest Mesoamerican allies.     

Somatic embryogenesis and organogenesis in Zamia fischeri,Z. furfuracea and Z. pumila

Morphogenesis from cultured megagametophyte, nucellus and immature embryos of three Zamia species on modified B5 medium containing 2,4-d and kinetin was compared. Organogenesis and somatic embryogenesis occurred from the megagamethophyte and zygotic embryo explants of Zamia pumila and Z. furfuracea tissue cultures, but only from the megamametophyte of Z. fischeri. Nucellar callus of Z. pumila produced globular structures that failed to develop further. Plantlets were recovered from somatic embryos of Z. pumila.Abbreviations 2,4-d 2,4-dichlorophenoxyacetic acid     

Branchinecta oterosanvicentei n. sp. (Branchiopoda: Anostraca), a new fairy shrimp from the Chihuahuan desert, with a proposal for the conservation of the Branchinectidae of Mexico

Branchinecta oterosanvicentei, new species, a fairy shrimp endemic to the south of Coahuila, Mexico is described and figured. Differential features between the new species and the related B. lindahli are discussed on the basis of a SEM micrographs. The main diagnostic characters of Branchinecta oterosanvicentei are: (1) a pulvinus covered by scales localized on the middle of the median side of the proximal article of antenna, and (2) a network of prominent cortical crests on the cyst surface. Of seven Branchinecta species occurring in Mexico, four (B. belki, B. mexicana, B. oterosanvicentei, and B. sandiegonensis) have a restricted geographic range. Because, in addition, extreme fluctuations in the number of mature individuals per population occur, the survival of all of these species is threatened, and measures should be taken to protect them. Branchinecta mackini has a wide distribution in North America, but even this form is rare in Mexico.     

A New Zamia Species from the Panama Canal Area

There are four terrestrial, above-ground stemmed Zamia taxa in Panama, the species delimitations of which have been a matter of controversy or misplacement at one time or the other (Schutzman et. al., 1998; Stevenson, 1993; Taylor, 1999a, b, 2002). All are allopatrically distributed. Two are in western Panama in Chiriquí province. One of these, Z. pseudomonticola, is found in the northwest of the province at altitudes above 1000 m, while the other, Z. fairchildiana, is found in a relatively small patch of forest in southwestern Chiriquí province. The other two taxa are found around the Canal area or farther east. The species described in this paper is found near the Canal area, and the last of the group, Z. elegantissima, occurs north of the Canal area in the province of Colon and also some distance to the east, including part of the Dule or Kuna aboriginal homeland known as Kuna Yala. After 16 years of research on the new taxon, we have decided to describe it as a new species, pointing out its similarity to Z. elegantissima and its distinctness from Z. pseudomonticola and Z . fairchildiana. The two western species are more alike in structure compared with the eastern species, and the latter are more alike between themselves compared to the western taxa. Even so, there are differences in vegetative and reproductive structures to clearly separate each species. There are even differences in the pollinators, these being, in all cases found, species of the weevil genus Rhopalotria and the snubbed-nosed beetle Pharaxonotha.     

Spathacanthus magdalenae sp. nov. (Acanthaceae), a riparian forest species from Veracruz,Mexico

Spathacanthus magdalenae Cast.‐Campos (Acanthaceae) is described and illustrated as a new riparian forest species from central Veracruz and from the eastern slopes of the volcano Cofre de Perote, Mexico. The new species is similar to Spathacanthus parviflorus, but differs in wider leaves and longer corolla tube. A key to all four species of Spathacanthus is provided.     

Zelandoperla maungatuaensis sp. n. (Plecoptera: Gripopterygidae), a new flightless stonefly species from Otago,New Zealand

ABSTRACT

A new wing-reduced species of the stonefly genus Zelandoperla Tillyard is described from Otago, New Zealand. Zelandoperla maungatuaensis sp. n. differs from a related species, Z. denticulata, primarily by lacking full wing development. We used morphological, geographical, and genetic evidence to assess the proposed status of Z. maungatuaensis sp. n. and Z. denticulata as separate monomorphic species, as opposed to conspecific members of a polymorphic species. High genetic divergence was found between distinctly allopatric populations, supporting the status of Z. maungatuaensis sp. n. as a distinct monomorphic species.     

<Emphasis Type="Italic">Agave manantlanicola</Emphasis> (Agavaceae), a new species from western Mexico

Agave manantlanicola, a new endemic species from the high altitude mountains of the Sierra de Manantlán in western Mexico, is described and illustrated. The species belongs to the subgenus Littaea and Gentry’s Amolae group. The new species is compared to other morphologically similar species from western Mexico, including Agave attenuata subsp. attenuata, A. attenuata subsp. dentata, A. bakeri, A. chazaroi, and A. vazquezgarciae. Information about its distribution, habitat, and conservation status is included.