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Hechtia kruseana and Hechtia rosamariae (Bromeliaceae; Hechtioideae), two new species from Guerrero, Mexico
Abstract and Figures
Recent botanical explorations carried out in the Guerrero state for the project Bromeliaceae of Mexico led to the discovery of two new species, both endemic to Guerrero: Hechtia kruseana and H. rosamariae. The new taxa are compared with the morphologically similar: H. anarosae, H. caulescens, H. jaliscana and H. stenopetala. Hechtia kruseana differs from these by the diameter of the rosettes (50-60 cm) and by the length of the staminate and pistillate inflorescences (190-275 cm). Hechtia rosa-mariae differs from morphologically related species by the size of floral bracts (1.5-2.0 × 0.7-1.0 mm) and the length of the petals (3.0-3.2 mm) of staminate flowers; by the length (2.3-2.6 mm) and shape (ovate) of the sepals, and width of the petals (1.7-2.0 mm) of pistillate flowers. Morphological descriptions, images, and a map showing the distribution of the described taxa are included, as well as an identification key for all the species of Hechtia present in Guerrero.
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The genus Hechtia includes 91 species (with those described herein), 88 of them present in Mexico, and the entire genus is endemic to the region termed “Megaméxico 3” by Rzedowski. With 21 species, the Mexican state of Puebla is the second most species rich. As a result of botanical exploration for the project Bromeliaceae of Mexico, we describe here seven new species, all endemic to Puebla: Hechtia anarosae, H. dasylirioides, H. espejoana, H. longissimifolia, H. microcarpa, H. minimiflora, and H. vicesphaeroblasta. The proposed species are compared with morphologically similar taxa (H. caulescens, H. ensifolia, H. flexilifolia, H. fragilis, H. podantha, and H. sphaeroblasta). Morphological descriptions, images, and a map with distributions of the described taxa are included, as well as an identification key and a list of specimens examined of all the species of Hechtia currently known from the state of Puebla.
The most recent phylogenetic analyses using molecular and morphological data of Hechtioideae revealed the presence of three well-supported, morphologically distinct clades related to each other as follows: (Hechtia tillandsioides complex [Hechtia guatemalensis complex (Hechtia s.s.)]). (1) H. tillandsioides complex is recognized here at the generic level (under the reestablished name Bakerantha), characterized by its grass-like leaves, which are almost entire or minutely dentate, central inflorescences with pedicellate flowers, and papyraceous, pendent fruits; this clade includes four species confined to the Veracruzan, Sierra Madre Oriental, Balsas Basin, and Transmexican Volcanic Belt biotic provinces. (2) The H. guatemalensis complex, here proposed as the new genus Mesoamerantha, is characterized by the presence of central inflorescences and flowers with 3/4 superior ovaries and is confined to the Pacific Lowlands, Veracruzan, Mosquito, and Chiapas Highlands provinces (in Belize, Guatemala, Honduras, El Salvador, and Nicaragua). The remaining sampled taxa are grouped into a clade (3) that consists of three well-supported lineages: the Hechtia glomerata complex distributed in the drainage of the Gulf of Mexico; a clade conformed by two species (H. deceptrix and H. epigyna) from the Sierra Madre Oriental that share an inferior ovary, and a poorly resolved internal clade (Core Hechtia) with the remaining species containing several well-supported, geographically restricted clades.
p> Background : Bromeliaceae family in Mexico has been the object of interest by botanists since 1789; their systematic study was approached from the 1970s onwards, and now there are significant advances in its taxonomic-floristic knowledge.
Question: How many and which species of Bromeliaceae occur in Mexico? How they are distributed, and how many are endemic?
Study site : México, 1887-2017.
Methods : Based on the study of the Mexican Bromeliaceae, including botanical collection, literature review, and revision, analysis and determination of specimens in 50 herbaria, data about species richness, Mexican endemics, and distribution of their taxa in the country, were obtained.
Results : In Mexico are represented four of the eight subfamilies of Bromeliaceae, 19 genera, 422 species, and 8 infraespecific taxa. The genera with the highest number of species in the country are Tillandsia (230/54.5 %), Hechtia (71/16.8 %) and Pitcairnia (50/11.8 %). 318 of the Bromeliaceae species are endemics to Mexico, as well as Ursulaea and Viridantha genera ; 172 species are microendemic. The entity with the highest number of taxa is Oaxaca, followed by Chiapas, Veracruz and Guerrero. Tlaxcala and Baja California Sur have the lowest species number. Baja California, Baja California Sur, Campeche, Ciudad de México, San Luis Potosí, Sonora, Tabasco y Tlaxcala have not strict endemic taxa.
Conclusion : Although progress in the knowledge of Mexican Bromeliaceae has been constant, exploration and recollection work is still required before concluding the Mexican bromeliad flora. It is also necessary to promote studies considering aspects of conservation and sustainable use.</p
We provide a map of the 14 biogeographic provinces of Mexico based on the ecoregions recognized for the country, which combine climatic, geological and biotic criteria. These provinces belong to the Nearctic region (Californian, Baja Californian, Sonoran, Chihuahuan Desert and Tamaulipas provinces), Neotropical region (Pacific Lowlands, Balsas Basin, Veracruzan and Yucatán Peninsula provinces) and the Mexican transition zone (Sierra Madre Occidental, Sierra Madre Oriental, Transmexican Volcanic Belt, Sierra Madre del Sur and Chiapas Highlands provinces). In order to facilitate future biogeographic analyses, we provide a file of the biogeographical regionalisation of Mexico by converting the map into a polygon shapefile and a raster file with all provinces. We also separately provide each of the provinces in vector and raster format. All the maps are in geographical and Lambert Conformal Conic projections.
Hechtia caulescens Lopez-Ferrari, Espejo & Martinez-Correa (Bromeliaceae), known from the states of Morelos, Puebla, and Oaxaca in Mexico, is described mid illustrated. The new species is compared with H. stenopetala Klotzsch, front which it differs by its larger flowers and floral bracts, and by its twice-branched inflorescences.
Cladistic analysis of ndhF sequences identifies eight major bromeliad clades arranged in ladderlike fashion. The traditional subfamilies Tillandsioideae and Bromelioideae are monophyletic, but Pitcair- nioideae are paraphyletic, requiring the description of four new subfamilies, recircumscription of Pit- cairnioideae and Navioideae, the sinking of Ayensua, and description of the new genus Sequencia. Brocchinioideae are basalmost, followed by Lindmanioideae, both restricted to the Guayana Shield. Next is an unresolved trichotomy involving Hechtioideae from Central America, Tillandsioideae, and the remaining bromeliads in subfamilies Navioideae, Pitcairnioideae, Puyoideae, and Bromelioideae. Bromeliads arose as C 3 terrestrial plants on moist infertile sites in the Guayana Shield roughly 70 Mya, spread centripetally in the New World, and reached tropical West Africa (Pitcairnia feliciana) via long-distance dispersal about 10 Mya. Modern lineages began to diverge from each other 19 Mya and invaded drier areas in Central and South America beginning 15 Mya, coincident with a major adaptive radiation involving the repeated evolution of epiphytism, CAM photosynthesis, impounding leaves, several features of leaf/trichome anatomy, and accelerated diversification at the generic level. This ''bromeliad revolution'' occurred after the uplift of the northern Andes and shift of the Amazon to its present course. Epiphytism may have accelerated speciation by increasing ability to colonize along the length of the Andes, while favoring the occupation of a cloud-forest landscape frequently dissected by drier valleys. Avian pollination (mainly by hummingbirds) evolved at least twice ca. 13 Mya; entomophily was ancestral. Hechtia, Abromeitiella-Deuterocohnia-Dyckia-Encholirium, and Puya exhibit a remarkable pattern of concerted convergence in six anatomical and physiological leaf traits adapted to drought.
El monto de la flora fanerogámica de México se calcula en forma aproximada en ±220 familias, ±2410 géneros y ±22000 especies. La mayor concentración de la diversidad se encuentra a lo largo de un área que se inicia en Chiapas, incluye Oaxaca, propagándose por un lado hacia el centro de Veracruz y por el otro a Sinaloa y Durango. El bosque mesófilo de montaña y el bosque tropical perennifolio son los más diversos por unidad de superficie; sin embargo, en números absolutos de especies quedan superados por otros tipos de vegetación. La proporción de taxa endémicos a los límites del país se aproxima a ±10% en el caso de los géneros y a ±52% en cuanto a las especies; estos valores aumentan a ±17% y a ±72% respectivamente si se toma como marco de referencia a un área ecológicamente más natural, pero que incrementa en ±1/3 la superficie de México. El endemismo se manifiesto más conspicuamente en la flora de matorrales xerófilos y de pastizales, aunque a nivel de especie también es intenso en otros tipos de vegetación, salvo el bosque tropical perennifolio. Las cifras anteriores son indicadoras de que el territorio del país ha sido de origen y de desarrollo de un gran número de linajes vegetales y en particular: En las zonas áridas y semiáridas del norte de México las plantas han sufrido una evolución profunda, dando origen a una flora moderadamente rica de sello propio y de formas biológicas especializadas, en no pocos casos únicos; La flora de las regiones semihúmedas se desarrolló en gran proporción a base de elementos que existen también en otras partes del mundo; un considerable número de tales elementos ha experimentado localmente una extensa radiación secundaria, produciendo una flora cuantiosa y diversificada; La flora de las regiones húmedas, sobre todo de las cálido-húmedas, sobre todo de las cálido-húmedas del este y del sureste del país también es muy variada, pero hasta ahora no hay muchos indicios de que México pudiera haber sido un centro importante de su evolución. El análisis de las unidades geográficas de la flora fanerogámica de México indica que su vinculación con el sur es unas cuatro veces más importante que con el norte. Tal hecho, sin embargo, no debe interpretarse en el sentido de que una mayoría tan grande de plantas mexicanas sea de derivación meridional directa, pues una buena parte de los elementos comunes con Centro y Sudamérica deben haberse originado en México o en otras partes del mundo, como las Antillas, África, Eurasia o Norteamérica. El registro fósil hasta ahora conocido indica que las características fundamentales de la actual flora fanerogámica de México estaban ya bien establecidas desde el Terciario Medio y muchas de ellas posiblemente desde tiempos anteriores.
El estado de Guerrero resulta de especial interés por la singularidad de su flora bromeliológica, la cual presenta un alto porcentaje de endemitas. Con base en el estudio de 1,250 ejemplares herborizados correspondientes a 797 recolecciones y una detallada revisión bibliográfica, concluimos que la flora bromeliológica hasta ahora conocida para Guerrero consta de 88 especies y 11 géneros. Se registran por primera vez los géneros Racinaea y Ursulaea y las especies Catopsis berteroniana, Hechtia reflexa, Racinaea ghiesbreghtii, Tillandsia andrieuxii, T. gymnobotrya, T. magnusiana, T. marabascoensis, T. polystachia, T. rettigiana, T. tricolor y Ursulaea tuitensis. Los estados con una flora bromeliológica más parecida a la de Guerrero son Jalisco, Michoacán y Colima, seguidos por el Estado de México y Morelos. De los tipos de vegetación que prosperan en el estado, el que presenta una mayor riqueza bromeliológica es el bosque de Quercus con 42 taxa, seguido por los bosques tropical caducifolio con 33 y tropical subcaducifolio con 27. El tipo de vegetación con menor número de especies es el bosque de Juniperus. Se presenta un catálogo completo de la familia, incluyendo claves de identificación a nivel genérico y específico, y se hace un análisis de la representación de la familia en el estado, documentándose detalladamente la distribución de sus especies.
The genus Hechtia includes 84 species, 79 of them present in Mexico. With 12 species (including those described herein), the Mexican state of Guerrero is the third most diverse in species. As a result of botanical explorations for the project Bromeliaceae of Mexico, we describe here five new species for science, all endemic to Guerrero: Hechtia elegans, H. ensifolia, H. medusae, H. platyphylla, and H. pycnostachya. The proposed species are compared with other morphologically similar taxa (H. caulescens, H. flexilifolia, H. glauca, H. hintoniana, H. laxissima, and H. pretiosa). Morphological descriptions, images, and a distribution map of the described taxa are included, as well as an identification key for all the species of Hechtia currently known from Guerrero
Se presenta un listado actualizado de las Liliopsida endémicas de México. De las 4,542 especies silvestres que habitan en el territorio nacional, 2,010 son endémicas exclusivas y la cifra asciende a 2,764 si consideramos la ampliación de los límites a lo que Rzedowski denomina Megaméxico 3. Se incluyen datos relativos al número de taxa endémicos por familia, por género y por estado, así como otros análisis relativos a la representación del endemismo por forma biológica y algunos ejemplos de endemismo ecológico.
Two new species are described in the genus Hechtia: H. carlsoniae and H. pumila. Hechtia melanocarpa, previously thought to be known only from its type, is represented by several collections. In contrast to the type specimen which is of horticultural origin, collections from the type locality have consistently larger inflorescences with elongate lateral branches. Moreover, staminate and pistillate individuals differ significantly in inflorescence form and petal shape.
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