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1 Sanguisuga, a New Genus of Neotropical Cytinaceae And http://www.icn.unal.edu.co/ Caldasia 34 (2): 291-308. 2012 SANGUISUGA, A NEW GENUS OF NEOTROPICAL CYTINACEAE AND SOUTH AMERICAN CONNECTION IN THE FAMILY Sanguisuga, Un Género Nuevo Neotropical De Cytinaceae Y Una Conexión Sudamericana En La Familia José Luis Fernández-Alonso Real Jardín Botánico, RJB-CSIC, Consejo Superior de Investigaciones Científicas, Plaza de Murillo 2, 28014 Madrid, España. [email protected]: Autor correspondencia. Hermes Cuadros-Villalobos Postgrado Biología-Línea Biodiveridad y Conservación, Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apartado 7495, Bogotá D.C. Colombia. ABSTRACT Sanguisuga caesarea, a new genus and species of Cytinaceae from Colombia and the first record of this family in South America, is described and illustrated. Sanguisuga is close to Bdallophytum but differs from it by exhibiting andromonoecia, with male flowers only at the base of the inflorescence, flowers zygomorphic, semiclosed, and dorsoventrally compressed. The tongue-shaped tepals are arched, curved inwards and include a ventral, sigmoid, nectar-secreting tepal. The elongated seeds are obpyriform with a length/width ratio of 1,5-1,8. Key words. Bdallophytum, Colombia, Cytinaceae, parasitic plants, South America, taxonomy. RESUMEN Se describe e ilustra a Sanguisuga caesarea, un género nuevo y una especie nueva de Cytinaceae de Colombia y el primer registro de esta familia en Sudamérica. El género nuevo es afín a Bdallophytum pero se diferencia por presentar andromonoecia, con las flores masculinas en la base de la inflorescencia, flores zigomorfas comprimidas dorsiventralmente, semicerradas, con los tépalos arqueados e incurvos; tépalo inferior- medio sigmoide-lingüiforme y nectarífero y las semillas más alargadas, obpiriformes con una relación largo/ancho de 1,5-1,8. Palabras clave. Bdallophytum, Colombia, Cytinaceae, plantas parásitas, Sudamérica, taxonomía. INTRODUCTION In the family Cytinaceae two genera, Cytinus L. and Bdallophytum Eichler, and a dozen species have so far been recognized. This small family of holoparasitic angiosperms, endophytes of roots, having flowers grouped in inflorescences and connate stamens, is considered in the current phylogenetic system as part of the order Malvales (Nickrent 2002, 2007, Nickrent et al. 2004, 1 Stevens 2011), and as a group independent of Rafflesiaceae s.l., a family that previously had been treated two genera (Harms 1935, Meijer 1993). Within Malvales, it is considered that the neotropical family Muntingiaceae (Bayer 2003), is the closest photosynthetic relative to Cytinaceae with which it shares some characteristics such as type of indumentum, pollen, position of ovary and placental type (Nickrent 2007). Cytinus includes at least 7-8 species parasitic on Cistaceae, Asteraceae and Rosaceae and is distributed mainly in the Mediterranean, South Africa and Madagascar (Burgoyne 2006, Vega et al. 2007, 2010 Grayum, Nickrent 2011). Bdallophytum known to have only three neotropical species, restricted to the region between Mexico and Costa Rica, with known records from Mexico, Guatemala, Nicaragua and Costa Rica (Eichler 1872, Kuijt 2001, Alvarado 2009 Grayum 2010). The species of this genus usually live associated with the roots of the genus Bursera Jacq. ex L. (Burseraceae). The genus Bdallophytum, (whose name derives from the Greek bdell = leech and phyton = plant) is characterized by a spicate inflorescence, flowers purple or purplish, campanulate or rotate perigone and seeds with an ornate testa, allowing one to easily separate it from the genus Cytinus that presents flowers grouped in aggregated racemes or glomeruli (or more rarely solitary flowers), tubular perigone of brighter colors (red, yellow or white) and smooth seeds (Solms-Lubach 1874, 1901, Bouman & Meijer 1994, Alvarado 2009). A recent floristic study conducted in dry forests of northern Colombia, revealed the presence of a previously unknown parasitic plant in South America, which appeared associated with the roots of trees of the genus Bursera and presented upon preliminary analysis some similarities with the aforementioned genus in Cytinaceae, Bdallophytum. As a result of the study of these samples, there is disclosed herein a new genus of Cytinaceae, which presents some morphological innovations not previously documented in this family. In turn, this finding represents an extension of the distributional range for Cytinaceae of about 800 km further south and confirms a hypothetical South American connection in the origin and distribution of this family of parasitic plants (Nickrent 2007). The nearest known populations of Cytinaceae (Costa Rica), are about 2500 km distant in a straight line from those now located in northern Colombia (Fig. 7). Materials and Methods The material studied comes from the field work developed by the second author in the dry forests of Valledupar, in foothills of the Sierra Nevada de Santa Marta (Dept. of Cesar, in the Caribbean region of Colombia), where one can track the plant for recollecting fruiting specimens. The specimens were preserved in 70% ethyl alcohol, without crushing, being properly stored in the liquid sample collection (antotecas) of the COL and MA herbaria. Some sections of the plant, made with a scalpel blade, and these have been preserved as dry and mounted specimens with photos in the general herbaria of both institutions. The observations were made with a Nikon SMZ645 Binocular microscope and detailed photographs were taken with a Canon camera attached to it. Palynological samples (for optical microscopy) were prepared in the Palynology Laboratory of the Institute of Natural Sciences, National University of Colombia, following the acetolysis technique of Erdtman (1986) and for observing the micropreps we used a trinocular Zeiss microscope, model Axiostar Plus. A Canon Power Shot A70 was used with the 100X lens for photographs. All measurements were taken in microns (µ) with ocular micrometer. The palynological slide collection was deposited in the palynological collection of ICN of the 2 Universidad Nacional de Colombia. For scanning electron microscopy (SEM) samples were mounted on aluminum supports, coated with a 40-50 nm layer of gold and examined with a Hitachi S-3000N scanning electron microscope at 15 kV. For the description of palynomorphs we followed the terminology of Punt et al. (1994). Descriptive terminology. For descriptive terminology was basically followed that recently used for this group (Alvarado 2009, Stevens 2011 and Nickrent 2007, 2011). However, we present here then some dimensions and morphological details of certain organs or structures of this plant, not listed as such in the literature. Haustorial nodules: In the roots parasitized by the endophyte are hemispherical or discoid protruberances, in mode of buds, with smooth surface (young) and generally less than the root diameter (Figs 1a, 3a). Floriferous Nodules: globose or conical structures, constituted by clusters of horny, linear or narrowly triangular scales. They occur along parasitized host roots and the shoots originate by tearing through the haustorial nodules (Figs 1a, 3b, d.) Caudex or haustorial root stock: it is a term used to refer to the scaly thickening accompanied by two or more floriferous nodules, which occasionally occur in the lattice, where two or more parasitized roots converge (Figs. 2a, 3a). Perigone: Used because for the petaloid perianth when not differentiated into calyx and corolla and apparently originated in a single whorl (calyx) of tepals (Figs. 1d, l, 4f, g). Coenocarp: We use this name because it is a multiple fruit type, constituted by both the fleshy axis of inflorescence and individual ovaries, partially or completely encompassed in the axial tissue (Fig. 4). RESULTS Sanguisuga caesarea Fern. Alonso & H. Cuadros gen. et sp. nov. Fig. 1 TIPO: COLOMBIA. Departamento del Cesar. Estribaciones de la Sierra Nevada de Santa Marta, Mun. de Vallepdupar, Corregimiento Rioseco, finca “Las Palomas”, 10º 40’ N, 73º 7’ E, 290 m, 6 Mar 2011, fl, H. Cuadros, D. Cortés & C. Delgado 6333 (holotype COL, isotypes COL, MA). Figs. 1-6. New genus and new species, similar to genus Bdallophytum Eichler (Cytinaceae) but differs by being a andromonoecious plant with hermaphroditic flowers, male flowers only at the base of the inflorescence and lacking female flowers; with zygomorphic flowers, dorsiventrally compressed, semiclosed, with tepaloid perigone, 7-9(10) laciniated; tepals unequal, arched, curved inwards, with a ventral, sigmoid, tongue-shaped, nectar-secreting tepal; seeds obpyriform, with a length/width ratio of 1,5-1,8. Description. Plant herbaceous, achlorophylous, holoparasitic and andromonoecious (hermaphrodite and female flowers on the same individual); endophyte vegetative tissue (not visible) developed in shallow roots of plants of the genus Bursera L. (Burseraceae). Parasitized roots generally horizontally disposed and with a rhizomatous aspect, located about 8.12 cm depth, of several decimeters in length and 3-4.5 (6) mm thick, with shiny, crust, fragmented and peelable (Figs. 1a, 2a); with smooth nodose formations similar to buds (haustorial nodules) 2.5-3 mm in diameter, irregularly distributed along the parasitized roots; with floriferous nodules (Figs 1a, 3b, d) dispersed in roots, globose or conical, rigid and with an echinate aspect, consisting of groups of horny scales, linear or narrowly triangular, occasionally with several floriferous nodules grouped at the confluence of several roots (Figs. 2a, 3a), forming 3
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