Biota Colombiana 8 (1) 1 - 45, 2007

Diversity of Colombian Passifloraceae: biogeography and an updated list for conservation

John Ocampo Prez1, Geo Coppens dEeckenbrugge2, Mara Restrepo1, Andy Jarvis1,3, Mike Salazar1, and Creuci Caetano1,4. 1 Bioversity International (formerly IPGRI), Regional Office for the Americas, A.A. 6713, Cali, Colombia. E-mail: [Link]@[Link] 2 CIRAD/FLHOR, UPR Gestion des ressources gntiques et dynamiques sociales, Campus CNRS/Cefe, 1919 route de Mende, 34 293 Montpellier, France. 3 International Center for Tropical Agriculture (CIAT), A.A. 6713, Cali, Colombia. 4 Universidad Nacional de Colombia Sede Palmira. Facultad de Ciencias Agropecuarias. Kra. 32 Chapinero, va Candelaria. Palmira, Valle del Cauca, Colombia.
Abstract The list of Colombian Passifloraceae was revised, using 3.930 records from literature, herbaria, and field observations. It includes 167 species, 165 of them native, which is equivalent to 27% of the family. Our list includes more details on species distribution and presents 26 species new to Colombia. Passiflora is the most important genus, with 162 species, whose center of diversity is in the Ecuadorian and Colombian Andes. Inside Colombia, the highest diversity is concentrated in the Andean region, which houses 81% of the species, particularly in the departments of Antioquia, Valle del Cauca, Cundinamarca, Quindo, Risaralda, and Caldas. The highest number of species is found at between 1000 and 2000 m above sea level and the most common thrive in disturbed habitats, such as roadsides, cultivated plots, and secondary forests. Most of the 58 endemic species are found at between 1500 to 2500 m and belong mainly to subgenera Tacsonia and Decaloba. Forty-two species produce an edible fruit, and nine are commercially cultivated. Among the species reported, 70% are threatened to some degree and three are considered extinct. Colombia may still house many unknown species in poorly explored departments, but more information about Passiflora diversity and distribution is needed to develop its economic potential. The conservation of this threatened species along with its habitat is essential and urgent. Because of the species multiple ecological interactions with many organisms, both aspects can be combined using Passifloraceae as an indicator of biodiversity in the Andean region. Keywords: biogeography, biodiversity, Colombia, Neotropics, Passifloraceae, passionflower, threatened species Resumen La lista de Passifloraceae colombianas fue revisada, usando 3.930 datos provenientes de la literatura, herbarios, y observaciones de campo. Incluye 167 especies, de las cuales 165 son nativas, representado el 27% de la familia. Nuestra lista trae ms detalles de la distribucin de las especies y presenta 26 especies nuevas para Colombia. Passiflora es el gnero ms importante, con 162 especies. En comparacin con otras regiones, los Andes de Colombia y del Ecuador constituyen su centro de la diversidad. Dentro de Colombia, la mayor diversidad se concentra en la regin andina con 81% de las especies, particularmente en los bosques de las cuencas hidrogrficas entre 1000 y 2000 m, en los departamentos de Antioquia, Valle del Cauca, Cundinamarca, Quindo, Risaralda, y Caldas. Las especies comunes crecen generalmente en habitats disturbados, como bordes de caminos y de cultivos, y bosques secundarios. La mayora de las especies endmicas (58) son encontradas entre los 1500 y 2500 m, y pertenecen principalmente a los subgneros Tacsonia y Decaloba. Veinte y dos especies producen un fruto comestible, y nueve se cultivan comercialmente. Entre las especies reportadas, 70% presentan algn grado de amenaza y tres se consideran extintas. Colombia puede ser el escenario de muchas especies desconocidas en departamentos poco explorados. Un mejor conocimiento de la diversidad del gnero Passiflora y de su distribucin es necesario para desarrollar su potencial econmico. Es una tarea urgente la conservacin de esta riqueza amenazada y de su habitat. Proponemos combinar ambos aspectos, utilizando las Passifloraceae como indicador de la biodiversidad en la regin andina, lo cual parece justificado por sus mltiples interacciones ecolgicas con otros organismos. Palabras claves: biogeografa, biodiversidad, Colombia, Neotrpico, Passifloraceae, flor de la pasin, especies amenazadas Biota Colombiana 8 (1), 2007

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Introduction he Passifloraceae consist of 18 genera and approximately 630 species, distributed throughout the tropics from the coastal zones up to 3800 m above sea level in the Andean paramos (Holm-Nielsen et al. 1988). In America, the family is represented by four genera (Ancistrothyrsus, Dilkea, Mitostemma and Passiflora), of which Passiflora, with about 530 species distributed mainly in the New World, is numerically and economically the most important genus of the family (Ulmer & MacDougal 2004). Only 22 species of the subgenus Decaloba (syn. Plectostemma sensu Killip) are distributed in the Old World, in the tropical and sub-tropical regions of Southeast Asia and Austral Pacific. Passionflowers are generally perennial lianas or herbaceous vines with tendrils, although some are trees, shrubs, or even annuals. Their wide morphological variation appears to result from the diversity of their habitats as well as their coevolutionary relationships with many organisms, including protective ants (Apple & Feener 2001), herbivores (particularly Heliconius spp. butterflies; Gilbert 1982), pollinators, and the plant communities providing them physical support and access to sunlight. Pollination is mainly carried out by insects and birds; several species are bat-pollinated (Endress 1994; Bchert & Mogens 2001), and a few species exhibit elements of the carnivory syndrome (Radhamani et al. 1995). Many species are cultivated for their edible fruit, as ornamentals, or for their medicinal properties (Ulmer & MacDougal 2004; Coppens dEeckenbrugge 2003; Martin & Nakasone 1970; Dharwan et al. 2004). P. edulis Sims (maracuja) is by far the best known and economically important species of the family. When Spanish missionaries arrived in South America in the 16th century, they felt that passionflowers were a good omen for their mission. In their unique morphology, they saw the elements of the Passion of Jesus Christ and a sign that the New World would successfully be converted to Christianity (Killip 1938; Uribe 1955a). This religious symbolism gave the plant its common name of Flos Passionis, or Passion Flower. The Latin translation by Pluckenet (1696) was accepted for the genus Passiflora created by Linnaeus in 1753, who described 24 species in his Species Plantarum, a number increased to 35 by Lamarck (1789). The first extensive monograph of the family was published by Cavanilles in 1780, with 43 species treated. They were followed by authors like Jussieu (1805), De Candolles (1828), Roemer (1846), Masters (1872), Triana & Planchon (1873) and Harms (1925), who described about 250 species divided into 21 sections (Killip 1938). In his 1938 monograph, The American Species of Passifloraceae, Killip made the most extensive description of the New World species, classifying 355 species into
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17 genera and 22 subgenera, based on floral morphology. In Colombia, the priest Uribe (1954, 1955a, 1955b, 1957, 1958, 1972) described several new species, and Escobar (1986, 1987, 1988a, 1988b, 1989, 1990, 1990 inedited, 1994) revised the subgenera Distephana, Manicata (syn. Granadillastrum), Rathea and Tacsonia, including Tacsoniopsis in the latter, and described one additional subgenus, Porphyropathanthus. She passed away in 1993, leaving an inedited document on her revision of subgenus Astrophea. MacDougal revised subgenus Plectostemma in 1994, restoring its ancient name Decaloba. In the last decade, MacDougal and Feuillet have published many papers including the description of about 15 new species, mainly of the subgenera Decaloba and Astrophea (MacDougal 1992, 1994, 2006; Feuillet 2002, 2004). Recently, Feuillet & MacDougal (2003) proposed a new infrageneric classification in Passiflora. According to this proposal, only based on morphological characters, four subgenera would be recognized: Astrophea and Deidamioides, from South and Central America; Decaloba, from America, Southeast Asia and Australia; and Passiflora, exclusively from America (Ulmer & MacDougal 2004). Additionally, they proposed to downgrade the genus Tetrastylis as a section of the subgenus Deidamioides. Recent molecular analyses (Muschner et al. 2003; Yockteng & Nadot 2004; Hansen et al. 2006) partly support the reduction in the number of subgenera, with the existence of at least three major groups, corresponding globally to subgenera Decaloba, Passiflora and Astrophea of the new proposal. On the other hand, molecular data from the different studies are not always consistent on the relative placement of these groups, and their results are less clear at lower levels, with inconsistent grouping of particular species and poor correspondence with some well established morphological divisions. In addition, the monophyly of Passiflora has not been established, and the study of Muschner et al. (2003) even raises some doubts about it. Clearly, more studies, involving more numerous species samples, are needed before reevaluating such a complex and fast evolving group as is that of the Passiflora. Colombia is the second most biodiverse country in the world (MacNeely et al. 1990). The country is divided into five main biogeographic regions: Amazon, Andes, Caribbean, Orinoco, and Pacific. The Andean region presents a highly varied topography (1000-5400 m) with three main mountain ranges. Thus, the Eastern, Central and Western Cordilleras separate two large inter-Andean valleys from the Pacific Coast to the West and the OrinoqueanLlanos to the East. The uplift of the Andes created new habitats and increased local isolation, favoring high speciation rates in many taxa. In Passiflora, a particularly striking example is given by subgenus Tacsonia, whose beautiful and large-flowered species are strictly adapted to high altitudes in cloud

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forests (2000-3800 m), and pollination by the sword-billed hummingbird Ensifera ensifera Lesson, which shows the same distribution (Bchert & Mogens 2001). As a result of this variety of habitats, Colombian flora includes one of the worlds most diverse groups of vascular plants, with 51.220 documented species (May 1992; UNEP-WCMC 2004). However, Colombia has undergone recent transformation of large parts of its natural ecosystems, in particular in the Andean region. Seventy percent of the Andes, an area vital to the conservation of Colombias water supply, has been deforested as a result of both agricultural colonization and human migration (World Press Review 1993). Destruction of natural habitats has drastically affected many species distributions, often reducing their historical ranges to a set of small, fragmented populations (Brooks et al. 2002). It has been predicted that such habitat alteration will lead to a substantial risk of extinction in the near future. Passifloraceae are of great interest within this context of rapid erosion of biodiversity, and not only for their fast radiation and spectacular variation in morphology and reproductive biology. Indeed, as stated above, this family is exemplative from the standpoint of coevolution in many respects, such as their particular relationship with specialized herbivores, ants and other nectar feeding insects; most importantly, they are parasites of structure, as they depend on many very different species for support, from low shrubs in disturbed habitats to high trees in primary forests. They are mainly perennials, but their life cycle is much shorter than that of their supports. They are sensitive to long-term changes in the ecosystem (dependence on trees) as well as short-to medium-term changes (by their other adaptive traits). Thus, they should constitute an excellent indicator group for the monitoring of biodiversity in Colombia. In addition, Colombia presents a long tradition of diversity in fruit production and consumption, and it is the country with the highest number of marketed passion fruit species, so the study of Passiflora diversity must also be thought of in terms of conservation of genetic resources of important or promising fruit crops. The last inventory by Hernndez & Bernal (2000) recorded 141 Passifloraceae species distributed in all the biogeographic regions. Forty-eight of them, mainly housed in the Andean region, are endemic to Colombia. This inventory was based on the study of specimens from five herbaria (COL, HUA, JAUM, MEDEL and MO), and the citations made in publications compiled by several authors that have worked on the family. Several recent collaborative projects have been focused on Passifloraceae. The Interamerican Development Bank (BID) has supported a regional project, coordinated by Bioversity International (formerly IPGRI) in 1994-1997.

Colciencias funded, in 1999-2001, the national project Conservacin y utilizacin de los recursos genticos de pasifloras, developed by French and Colombian scientists at the Bioversity Americas office. In 2004, the same group developed a study of diversity of the Passifloraceae and Caricaceae in the Colombian coffee growing area. All these projects have generated a considerable amount of information on morphology, cytology, palynology, molecular diversity, and biogeography of Passiflora, providing most of the material for the present inventory and allowing us to supplement and update the list of Hernndez & Bernal (2000) with new information, such as species new to science or to the country and elements of ethnobotanical information. In addition, the use of a Geographic Information System (GIS) allowed us to re-assess the conservation status of Colombian Passifloraceae species. Materials and methods Study Area Colombia is situated in the north of South America, between 12 26 46 N and 4 13 30 S, and between 66 50 54 W and 79 02 33 W, covering an area of 1,141,748 km2, with an altitudinal range from sea level up to 5775 m ([Link] The main administrative division defines 32 departments, and geographers recognize five biogeographic regions (Hernndez et al. 1991). Herbarium and Literature Data The data set consists of information gathered from specimen labels from 18 Colombian herbaria (AFP, CAUP, CDMB, CHOCO, COL, COAH, CUVC, FAUC, FMB, HUA, HUQ, JAUM, MEDEL, PSO, SURCO, TOLI, VALLE, UIS), and five herbaria in other countries (K, MA, MO, NY, P). These collections were gathered between 1750 and 2006. Most specimens were verified or identified, using the keys and descriptions of Killip (1938), Holm-Nielsen et al. (1988), Escobar (1988a, 1994), MacDougal (1994) and Tillet (2003). A synonymy list, based on the general list of Feuillet & MacDougal (2003), is given in Appendix 1. When possible, voucher label information was used to assign geographic coordinates to specimens, using gazetteers and topographic maps of Colombia (scale 1:50,000 and 1:250,000). The database was supplemented with materials mentioned in species descriptions, essentially those of Killip (1938, 1960), Uribe (1955a), and Escobar (1988a,b, 1989, 1990, 1990 inedited, 1994). Collection records with obviously inaccurate or doubtful data were excluded from the analysis. Coordinates were further checked by plotting all species on a dot map, using the DIVA-GIS 5.2 software (Hijmans et al. 2001). Finally, we followed the infrageneric classification by Killip (1938) with the amendments of Escobar (1988, 1989) and MacDougal (1994).
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Expeditions and Samples Collected The dot map of all geo-referenced specimens was used to plan germplasm collecting trips. The prioritization of explored areas followed three criteria: permission to access (unfortunately not obtained for protected areas), richness of species and collection gaps. The collecting trips were carried out during 2003-2006, covering 555 localities in 17 departments, between 0 and 4200 m of altitude. The explorations were concentrated in the Andean region, in watersheds, wild forest areas, cultivated fields and road edges. Data were recorded for each collected specimen, including locality names, elevation, geographic coordinates using a hand-held GPS device, status (wild, cultivated or introduced), and ethnobotanical information (if any). These passport data were recorded and tabulated. Finally, the Geographic Information System software DIVA-GIS 5.2 was used to generate a dot map of the distribution of accessions collected / observed during the expedition. Threat Status of Passifloraceae The distribution area of each native species was characterized by the maximum distance (MaxD) and the circular area (CA50), following the method of Hijmans et al. (2001). This methodology has been applied in a number of studies to provide quantitative assessment of the distribution area required by the Red List criteria, for example by Maxted et al. (2005). MaxD is the largest distance between any pair of observations of one species. CA50 is the total surface within a 50-km radius around all the observations for a same species. These methods were supplemented with historical records of each taxon and subjected to the Red List criteria of the World Conservation Union (IUCN 2003, 2004), involving complex combinations of quantitative observations concerning the size and structure of the population, the range and fragmentation of its distribution (extent of occurrence and area of occupancy), as well as the intensity of their past or foreseeable variation. Along these lines, we considered that CA50 under 20.000 km2, MaxD under 100 km and number of observations under six, as well as the absence of records younger than 100 years, are critical.

Data collecting A total of 3330 herbaria and 45 literature data, concerning 120 species, were gathered and georeferenced when coordinates were not directly available. The highest number of species and specimens were found in the Colombian herbaria COL and HUA, with 1056 and 976 records respectively. During the collecting trips, most specimens were observed in forest fragments, gallery forest and forest and road edges, mainly in the watersheds of the coffee growing zone, between 1000 and 2000 m. In all sites visited during the expeditions, 87 Passifloraceae species were recorded, of which five individuals could not be identified. The dot map in Figure 1 shows the spatial distribution of our final dataset of 3930 records per herbarium (3330), literature (45) and field collections (555) of Passifloraceae in the different biogeographic regions. Distribution of Species Richness The number of observations and species richness was highest on the Andean slopes with 123 species, followed by the Amazonian region with 45 species (Box 1). The Orinoquean region was the poorest, with only 18 species. The Andean and Caribbean regions share the highest number of species (27). By contrast, the Pacific and Caribbean regions only present four species in common. Figure 2 gives a synthetic image of the similarities in species occurrence among regions, confirming a relative similarity between the Amazonian and Orinoquean, as well as between the Andean and Caribbean regions. The Pacific Coast Passifloraceae appears relatively divergent. The Andean region, as well as the departments of Antioquia, Valle del Cauca, Cundinamarca and Santander displayed the highest richness of specimens and species (Box 2). Considering their area, Quindo, Risaralda and Caldas are even more diverse. The department of San Andrs and Providencia (Caribbean islands) are only represented by P. biflora Lam. and P. pallida L.

Box 1 Distribution of Passifloraceae by biogeographic region. The diagonal gives their contribution in species number (bold) and contribution to the countrys total. The other cells give the number and proportion of shared species for each pair of regions. Biogeographic region Amazonian Andean Caribbean Orinoquian Pacific amz 45 (28%) and 21 (14%) 123 (76%) car 9 (12%) 27 (20%) 38 (23%) ori 15 (31%) 7 (5%) 9 (19%) 19 (12%) pac 15 (23%) 14 (10%) 4 (6%) 9 (14%) 36 (22%)

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Figure 1. Map of distribution of Passifloraceae specimens for 3,930 collections on five biogeographic regions in Colombia. Points on the maps represent sites of collection.

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Box 2 Number of observations and species of Passifloraceae in the 32 Colombian departments. Department Amazonas Antioquia Arauca Atlntico Bolvar Boyac Caldas Caquet Casanare Cauca Cesar Choc Crdoba Cundinamarca Guaina Guaviare Huila La Guajira Magdalena Meta Nario Norte de Santander Putumayo Quindo Risaralda San Andrs y Providencia Santander Sucre Tolima Valle del Cauca Vaups Vichada Abbreviation ama ant ara at bl by cl cq cs cau ce cho cor cun gn gv hu lg ma met na ns pu qu ri sp snt suc to vc va vch Biogeographic region amz and car pac and ori car and car and ori and amz and and ori amz and pac and car and pac and car and ori amz amz and and car car amz and ori and pac and amz and and and pac car and car and and pac amz ori Observation number 87 784 10 18 33 145 245 47 4 161 13 211 33 419 16 27 62 21 71 85 170 79 56 150 68 4 203 6 213 420 35 16 Species number 19 70 6 7 17 36 36 18 4 42 10 40 9 53 10 14 22 12 31 24 44 36 26 38 24 2 48 3 44 56 20 9

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Figure 2. Diagram comparing the similarity in contribution of Passifloraceae species to the floras of the Colombian biogeographic regions (Jaccard distance).

New Passifloraceae Checklist for Colombia Box 3 gives the number of species for each genus and subgenus present in Colombia in relation with the number of species present in the Neotropics. The updated inventory of the Colombian species (Box 4) includes a total of 167 Passifloraceae species, representing three genera, Ancistrothyrsus, Dilkea and Passiflora. This is equivalent to 27% of all Passifloraceae. The genus Passiflora is by far the most important with 162 species, representing 11 of Killips subgenera, and all the four subgenera defined in the classification proposed by Feuillet and MacDougal (2003). The most abundant species were P. vitifolia Kunth (359 specimens) and P. mixta L. (162 specimens), while 67 species (23%) were represented by a single specimen. In the expeditions, we found some species that had not been collected in the last decades, such as P. erytrophylla Mast., P. guazumaefolia Juss., and the semi-arborescent P. mariquitensis Mutis ex Uribe. The latter was described in 1783 by Jos Celestino Mutis during the Botanical Expedition of the Nuevo Reino de Granada in Mariquita (Tolima). It was considered extinct by Uribe (1955a) and a synonym of P. pittieri Mast. by Escobar (1990 inedited). However, we could verify that P. mariquitensis still exists, as three specimens that we have collected in a forest with high distribution near Mariquita corresponded closely to the type specimen, while they appeared morphologically distinct from P. pittieri specimens from Costa Rica, Panama, and northwestern Colombia in several traits (e.g. nectar shape, peduncle length, nerve shape). Similarly, after comparing the collected materials with the type specimens, we maintained other species that had been considered synonyms by Hernndez & Bernal (2000), such as P. mollis Kunth H.B.K. (vs. P. cuspidifolia Harms), and P. hahnii (Fourn) Mast. (vs. P. guatemalensis S. Watson). Our list includes 26 species new to Colombia, from those recognized by

Killip (1960), Feuillet & MacDougal (2003) and Ulmer & McDougal (2004) and three inedited from Escobar (1990) and Hernndez (2003): Ancistrothyrsus antioquiensis L.K Escobar (ined.), P. alata Curtis, P. andina Killip, P. bucaramangensis Killip, P. candollei Tr. & Planch., P. chocoensis Gerlach & Ulmer, P. cincinnata Mast., P. hahnii, P. hirtiflora Jrgensen & Holm-Nielsen, P. killipiana Cuatrecasas, P. lyra Planch. & Linden & ex Killip, P. megacoriacea Porter-Utley (ined.), P. mollis, P. monadelpha Jrgensen & Holm-Nielsen, P. munchiquensis Hernndez (ined.), P. occidentalis Hernndez (ined.), P. pallida L. (clearly separated from P. suberosa by Porter-Utley, 2003), P. pillosissima Killip, P. popenovii Killip, P. sodiroi Harms, P. tuberosa Jacq., P. rigidifolia Killip, P. tricuspis Mast., P. truxillensis Planch. & Lind. P. caerulea L., recently introduced from Brazil and Argentina and cultivated as an ornamental, was not included in the counts of each department. P. alata was not counted for Quindo and Valle del Cauca either, as the material under cultivation was also introduced from Brazil. P. micrantha Killip was not included because Hernndez (2003) considered it a synonym of P. erythrophylla. Nine more species occur close to the Colombian international border (less than 100km), and possibly exist also in the country, although they have not been included in this inventory. Another important result is the presence of the genera Ancistrothyrsus and Dilkea in the Andean and Pacific regions, the former following the mention of A. antioquiensis by Escobar (1990 ined.), who, unfortunately, passed away before publishing her monograph on arborescent Passifloraceae. Several botanical forms and varieties are mentioned for P. edulis Sims, P. cumbalensis (Karst.) Harms, P. foetida L, P. ligularis Juss., P. longipes Juss., P. rugosa (Mast.) and P. tripartita (Juss.) Poir. A total of 42 species with edible fruit are reported. Nine of them are sold on the international, national and/or local markets, P. edulis f. flavicarpa Degener and P. edulis f. edulis (introduced), P. ligularis, P. tripartita var. mollissima, P. tarminiana Coppens & Barney, P. quadrangularis L., P. maliformis L., P. popenovii Killip, P. nitida Kunth, and P. alata Curtis. Other species, such as P. antioquiensis H. Karst., P. cumbalensis, P. laurifolia L., P. nitida Kunth, P. palenquensis Holm-Niels. & Lawesson P. tiliifolia L., and P. pinnatistipula Cav. are cultivated in home gardens. Some commonly cultivated species seem to depend on human activity for their propagation, which suggests an advanced stage of domestication and/or an incomplete acclimatisation following an ancient introduction. Thus, P. edulis f. flavicarpa, P. ligularis, P. quadrangularis L., P. popenovii, P. tripartita var. mollissima, and P. tarminiana, are exceptionally found as feral plants. The latter has pullulated as an invasive plant in Hawaii and New Zealand. Another particular
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case is P. edulis f. edulis, introduced from southern South America, which has naturalized at intermediate to high altitudes, where it is not uncommon in the wild. The vernacular names are very diverse for each species. In the Amazonian region, we noted several indigenous names for the species P. foetida var. gossypiifolia Desv. (Iana-leeg, Murulale), P. holtii Killip (Guachique), P. nitida (Burucua, Gemarundare, Tuchica, Jino-Goj), P. serratodigitata L. (Cipo-Cipo), P. vitifolia (Maloca de Fisi). In the Cauca and Nario departments (south of the Andean region) P. fimbriatistipula Harms and P. liguBox 3 Number of Passifloraceae species in Colombia and the Neotropics. Genus Ancystrothyrsus Dilkea Mitostemma Passiflora Astrophea Decaloba Dysosmia Distephana Manicata Passiflora Porphyropathanthus Psilanthus Rathea Tacsonia Tryphostemmatoides All Passifloraceae Subgenus

laris are named Pachuaca and Awapit in the indigenous languages. Among the species collected in our expeditions, we found several species growing very commonly in disturbed habitats like the road edges, secondary forest margins, and especially riverbanks between 1000 and 2000 m: P. adenopoda Moc, & Sess ex DC., P. alnifolia Kunth, P. coriaceae Juss., P. capsularis L., P. rubra L, and P. suberosa L. The latter two are considered weeds in the coffee plantations. At higher altitudes (above 2500 m), P. mixta is also very common in disturbed habitats.

Colombia 2 3 0 22 52 2 6 1 38 1 3 2 30 4 167

Neotropics 3 5 3 57 190 20 15 5 156 1 4 3 55 7 533

Endemism Among the 165 native species, 58 (36%) are endemic to the country. The largest concentration of these occurs in the Andean region, principally in the Cordillera Central, in the departments of Antioquia and Tolima. The elevation belt between 1500 and 2500 m presents the highest richness of endemic and rare species ( 5 observations). Only eight of these were represented with only one specimen (e.g. P. cremastantha Harms), while P. bogotensis Benth and P. antioquiensis were the most common endeBiota Colombiana 8 (1), 2007

mic species, with 23 recorded specimens each. The proportion of endemic species varied considerably among taxonomic groups, especially among the subgenera of Passiflora (Box 4). Thus, Tacsonia (21), Decaloba (14), Passiflora (9) and Astrophea (7) present the highest number of endemic species. Subgenus Tacsonia displays the highest richness of endemic species in the Cordillera Central with eight species, mainly of the Colombian section characterized by a very long peduncle (P. flexipes Triana & Planch., P. linearistipula L.K. Escobar,

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P. quindiensis Killip and P. tenerifensis L.K. Escobar). Twenty-one species (37%) are restricted to very small areas of one department. These are located mainly in the departments of Antioquia (7), Tolima (4), Santander (3), Cauca (2), while only one such narrow endemic species is found for the departments of Bolivar, Boyac, Choc, Caldas, Cauca, and Magdalena. Threatened Species The distribution parameters of the 165 Colombian Passifloraceae native species are given in Appendix 2, and Figure 3 shows their repartition according to their threat status under the criteria of the IUCN (2003, 2004). Seventy-one percent of them are under some degree of threat, 10% being critically endangered (CR), 6.1% vulnerable (VU) or endangered (EN). Four of the 16 critically endangered species are endemic. All three extinct species (EX) belong to the Andean subgenus Tacsonia. Unfortunately, the only two species of genus Ancistrothyrsus are included in the category CR. Only 16% of the species were placed in the two categories LC and NT, least concern and near threatened. The species P. alata, P. megacoriacea Porter-Utley and P. rigidifolia Killip are placed in the DD category because of deficient data. The 29.3% classified in least concern, belong mostly to subgenera Decaloba and Passiflora with 18 and 14 species, respectively.

Figure 3. Percentual number of the threat status of 165 Passifloraceae native species under the IUCN criteria.

Box 4 List of 167 Passifloraceae species of Colombia. Fifty-eight endemic species are marked by an asterisk (*); twenty-six species new to Colombia by the abbreviation nr; nine species probably present in the country are indicated between square brackets. New records, for a given biogeographic region, department (abbreviated as in Tables 1 and 2) or elevation-range are indicated by bold letters. Abbreviations in bold letters in the Notes column correspond to the plant habits: shrub (Ab), tree (Ar), and climber (Tr). V.N and I.N. indicate vernacular and indigenous names, respectively. Taxon Genus Ancistrothyrsus Harms, 1931 Ancistrothyrsus antioquiensis L.K Escobar (ined), 1988 * nr Escobar & Roldn 8819 (HUA) - Type F.J Roldn (pers. com.), Escobar (1990 inedited) Biogeographic Region Geopolitical Distribution Elevation Collection for Reference Bibliographic Reference IUCN Category Notes

and

ant

90-800

CR

Tr

[Ancistrothyrsus hirtellus A.H. Gentry, 1992]

amz

150-350

Gentry & Stein 47114 (MO) Isotype Vester & Matapi 639 (COAH)

Gentry 1992

Tr Reported in the Ecuadorian, Peruvian and Venezuelan Amazon. CR Tr

Ancistrothyrsus tessmannii Harms, 1931

amz

ama pu

50-400

Holm-Nielsen et al. 1988

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Colombian Passifloraceae

Ocampo et al.

Taxon Genus Dilkea Mast., 1871 Dilkea johannesii Barb. Rodr., 1885

Biogeographic Region

Geopolitical Distribution

Elevation

Collection for Reference

Bibliographic Reference

IUCN Category

Notes

amz

va

100-500

Soejarto 2461 (HUA)

Killip 1938

CR

Tr Tr V.N.: Canilla de Tente, Tripa de Tente (ama). Edible fruit

Dilkea parviflora Killip, 1938

amz

ama cq va

100-500

Gentry 64981 (MO)

Holm-Nielsen et al. 1988

LC

Dilkea retusa Mast., 1871

amz and pac

ama ant cho cq gv met pu snt va vc

100-500

Lpez et al. 5947 (COAH)

Killip 1938; Uribe 1955b; Holm-Nielsen 1974; HolmNielsen et al.1988

LC

Tr

Genus Passiflora L., 1753 Subgenus Astrophea (DC.) Masters, 1871 Section Astrophea Passiflora callistema L.K. Escobar, 1994 * Section Botryastrophea Passiflora holtii Killip, 1938 Jaramillo 7890 (COL) Killip 1938; Escobar 1990 Ind., 1994 Killip 1938; Holm-Nielsen et al.1988; Escobar 1990 Ind., 1994 Killip 1960; Escobar 1990 Ind., 1994 Killip 1938; Holm-Nielsen et al. 1988; Escobar 1990 Ind, 1994 Killip 1938; Holm-Nielsen 1974; HolmNielsen et al.1988; Escobar 1990 Ind., 1994 Tr I.N: Guachique, Bejuco (ama). Edible fruit car bl 100 E. Forero 487 (COL) - Type Escobar 1990 Ind., 1994 CR Tr Known only from the type.

amz

ama cq gn va

150-500

LC/NT

Passiflora pyrrhantha Harms, 1926

amz

va

400-1000

Shultes & Cabrera 12693 (COL) Betancourt et al. 9753 (COAH)

EN/CR

Tr

Passiflora securiclata Mast., 1893

amz ori

ara by gv va vch

150-500

LC

Tr

Passiflora spicata Mast., 1872

amz

gv

150-500

Cuatrecasas 7397 (COL)

VU

Tr

Passiflora spinosa (Poepp. & Endl.) Mast., 1871

amz and ori car

ama ant by cq cor cun gn met pu snt va vch

150-500

Zarucchi 4279 (COL)

VU

Tr V.N.: Cocorella (bl), Bejuco campano (snt)

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Ocampo et al.

Colombian Passifloraceae

-11

Taxon Section Dolichostemma Passiflora citrifolia (Juss.) Mast., 1871 Passiflora haughtii Killip, 1938 * Passiflora mariquitensis Mutis ex Uribe, 1954 * Passiflora mutisii Killip, 1938 *

Biogeographic Region

Geopolitical Distribution

Elevation

Collection for Reference

Bibliographic Reference

IUCN Category

Notes

amz

va vch

85-500

Barbosa & Zurucchi 2989 (COAH) Haught 1635 (COL) Ocampo et al. 55 (TOLI) Mutis 2279 (MA) - Type Gentry & Aguirre 15318 (COL)

Killip 1838; Escobar 1990 Ind. Killip 1938; Escobar 1990 Ind., 1994 Killip 1938; Escobar 1990 Ind., 1994 Killip 1938; Escobar 1990 Ind., 1994 Killip 1938; Escobar 1990 Ind., 1994; Gentry 1976

LC

Tr

and

snt

100-700

CR

Ab Ab Formerly considered extinct. Tr

and

to

420-700

CR

and

to

600

EX

Passiflora pittieri Mast., 1897 Section Euastrophea

pac

ant cho

50-1000

VU

Ab

Passiflora arborea Spreng., 1826

and car

ant bl by cau cl cun hu ma na qu ri to vc

1000-2300

Humboldt & Bonpland 5864 (P) - Type

Killip 1938; Prez 1956; Holm-Nielsen et al. 1988; Escobar 1990 Ind. Escobar 1994 Killip 1938; Escobar 1990 Ind., 1994 Killip 1938; Escobar 1990 Ind., 1994 Killip 1938; Holm-Nielsen et al. 1988; Escobar 1990 Ind., 1994 Killip 1938; Escobar 1990 Ind., 1994 Killip 1938; Uribe 1972; Escobar 1990 Ind., 1994

NT

Ar V.N: Cherimoyo (vc), Granadillo arboreo (cun). Edible fruit Ab

Passiflora lindeniana Planch. ex Triana & Planch., 1873 Passiflora emarginata Humb. & Bonpl., 1813 * Passiflora engleriana Harms, 1894 *

and

cun ns snt

1000-2700

Linden 1409 (P) - Type Humboldt & Bonpland (P) - Type Escobar 8853 (COL)

NT

and pac

cau cl cho na vc

1500-2000

LC

Ar Edible fruit

and

ant

1500-2500

VU/EN

Ar

Passiflora macrophylla Spruce ex Mast., 1883

amz and pac

ant cau cho pu na

60-1800

Alczar & Salgado 1203 (CAUP)

LC

Ab V.N: Acaba familia (cho)

Passiflora putumayensis Killip, 1938

and

pu

1350-2500

Mora-0. 3438 (PSO)

EN/CR

Ab Ar V.N: Gulupo de Arbol (cun), Capafraile (to). Edible fruit

Passiflora sphaerocarpa Triana & Planch., 1873 *

and

ant ce cun na ns qu ri snt to vc

400-1700

Schlim 285 (P) - Type

LC/NT

Biota Colombiana 8 (1), 2007

12-

Colombian Passifloraceae

Ocampo et al.

Taxon Passiflora tica GomezLaur. & L.D. Gmez, 1981 Section Pseudoastrophea

Biogeographic Region pac

Geopolitical Distribution ant cho

Elevation

Collection for Reference Escobar 2192 (HUA)

Bibliographic Reference Escobar 1990 Ind., 1994

IUCN Category LC/NT Ar

Notes

450-1500

[Passiflora costata Mast., 1872]

amz

50-350

Spruce 1670 (K) - Type

Killip 1938; Escobar 1990 Ind., 1994

Tr Reported in the Amazon of Peru, Brazil, Guianas, and Venezuela) (confluence of the rivers Rio Negro and Casiquiare). EN/CR Ar Tr Reported in the Amazon of Venezuela.

Passiflora grandis Killip, 1938 * [Passiflora ovata Martin ex DC., 1828]

and

ns snt

1000-2000

Schlim 585 (K) Colector n.v.

Escobar 1990 Ind., 1994 Killip 1938; Escobar 1990 Ind., 1994 Killip 1938; Holm-Nielsen 1974; Escobar 1990 Ind., 1994 Killip 1938; Escobar 1990 Ind. Killip 1938; Escobar 1990 Ind., 1994 Killip 1938; Escobar 1990 Ind.

ori

0-150

Passiflora phaeocaula Killip, 1927

amz ori

gn va vch

150-1100

Madrian 893 (MO,GH)

LC/NT

Tr Ab

Passiflora skiantha Huber, 1960

amz

gv

150-500

Cuatrecasas 7366 (COL) Tessmann 4385 (N) - Type Juncosa s.n. (JAUM) n.v.

NT/VU

Tr Tr Reported in the northern Amazon of Peru.

[Passiflora tessmannii Harms, 1926]

amz

50-500

Passiflora venosa Rusby Subgenus Decaloba (DC.) Rchb., 1828 Section Cieca Passiflora apoda Harms, 1929

and pac

cho

50-450

VU/EN

Tr

and

ant cau cl qu na ri to vc ant by cau cl cho cun hu ma ns qu ri snt to vc at bl ce

1900-3260

Hazen 9688 (MO) - Isotype Uribe 2565 (COL) Cuadros-H 1882 (COL)

Killip 1938; Hernndez 2003 Croat 1978; Holm-Nielsen et al. 1988 Killip 1938

LC/NT

Tr

Passiflora coriacea Juss., 1805 Passiflora holosericea L., 1753

and car pac

250-1500

LC

Tr V.N.: Ala de Murcielago

car

0-1400

LC/NT

Tr

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Ocampo et al.

Colombian Passifloraceae

-13

Taxon Passiflora megacoriacea Porter-Utley, 2003 nr Passiflora pallida L., 1753 nr Passiflora sodiroi Harms, 1922 nr Passiflora suberosa L., 1753 Section Decaloba Series Auriculatae

Biogeographic Region car

Geopolitical Distribution bl

Elevation

Collection for Reference Killip & Smith 14415 (US) Dugand & Jaramillo 2844 (COL) Escobar et al. 4368 (PSO) Cuatrecasas 15930 (VALLE)

Bibliographic Reference Porter-Utley 2003 Porter-Utley 2003 Holm-Nielsen et al. 1988 Holm-Nielsen et al. 1988

IUCN Category DD Tr

Notes

100-200

car

at bl ma sp

0-200

LC

Tr Appel Monkey (sp) Tr Tr V.N.: Curubita de Monte (ant)

and

cau ant cau cl cun gv na ns qu snt suc to vc

1850-2150

EN/CR

and car

200-2200

LC

Passiflora auriculata Kunth, 1817

amz and car ori pac

ama ant bl by cau cl cho cor cq cun gn gv met na ns pu qu snt to va vc vch

0-1500

Killip & Cuatrecasas 58988 (VALLE)

Holm-Nielsen 1974; HolmNielsen et al. 1988

LC

Tr V.N: Rejito (cun)

Series Sexflorae Passiflora sexflora Juss., 1805 Series Luteae Passiflora filipes Benth., 1843 Series Miserae Passiflora misera Kunth, 1817 Passiflora tricuspis Mast., 1872 nr ant at ara bl by cau cl cho cor cun cs lg ma met vc ns met E. Forero 9936 (COL) Estrada et al. 146 (MA) and qu ri vc 950-1250 Silverstone 7205 (CUCV) Holm-Nielsen et al. 1988 VU Tr and ant hu qu to vc 1700-2300 Zurucchi et al. 5813 (CHOCO) Holm-Nielsen et al. 1988 NT/VU Tr V.N.: Corvejo (na)

and car ori pac

0-1050

Killip 1938

LC

Tr

and

1220-2000

Killip 1938

CR

Tr Tr Reported in the Amazon of Ecuador, Brazil and Peru. Ornamental (qu)

[Passifora trifasciata Lemaire, 1868]

amz

ama pu

130-1100

Brandbyge et al.33556 (AAU)

Killip 1938; Nielsen et al. 1988

Series Punctatae Passiflora alnifolia Kunth, 1817 and car ant by cau cl cun ma na pu qu ri snt to vc 1400-2500 Hno. Daniel 2803 (MEDEL) Holm-Nielsen et al. 1988 LC Tr

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14-

Colombian Passifloraceae

Ocampo et al.

Taxon Passiflora andreana Mast., 1883 Passiflora azeroana L. Uribe, 1955 * Passiflora biflora Lam., 1789

Biogeographic Region and

Geopolitical Distribution ant cau cun ma na qu ri by cun hu snt ant at bl ce cl cho cun hu ma met na ns ri sp snt to vc by cun hu lg ma ns snt vc

Elevation

Collection for Reference GarciaB.12949 (COL) Lozano 3718 (COL) Garcia-B. 11720 (COL)

Bibliographic Reference Holm-Nielsen et al. 1988 Uribe 1957 Killip1938; Holm-Nielsen 1974; Croat 1978 Killip 1938; Holm-Nielsen 1974 Killip 1930, 1938

IUCN Category CR Tr

Notes

1500-3150

and

2500-3000

NT/VU

Tr Tr V.N.: Peyen Papaya (sp), Desjarretadera (cun) Tr V.N.: Curubo macho (cun)

and car

0-1500

LC

Passiflora bogotensis Benth., 1845 *

and car

2000-3700

Garcia-B. 15291 (COL) Killip & Smith 16787 (MO) Isotype Rudas et al. 2180 (COL) Cuatrecasas 12526 (COL)

LC

Passiflora bucaramangensis Killip, 1930 * nr Passiflora candollei Tr. & Planch., 1873 nr Passiflora chelidonea Mast., 1979

and

snt

1500-2600

EN

Tr

amz

ama ant ara cau cho na ns pu ri snt vc ant by cho cun ma met ns snt vc by cun snt

100

Killip 1938 Holm-Nielsen et al. 1988 Killip 1938; Hno. Daniel 1968; HolmNielsen 1974 Holm-Nielsen et al. 1988 Killip 1930, 1938; Hernndez 2003 Killip 1938; Uribe 1955a; Hernndez 2003 Killip 1938 Killip 1938; Prez 1956 Holm-Nielsen et al. 1988 Killip 1938; Hno. Daniel 1968

NT

Tr

and car pac

900-3000

LC

Tr

Passiflora cuneata Willd., 1809 Passiflora cuspidifolia Harms, 1893 Passiflora dawei Killip, 1930 * Passiflora erytrophylla Mast., 1872 * Passiflora lyra Planch. & Lind. ex Killip, 1846 nr Passiflora magdalenae Triana & Planch., 1873 * Passiflora micropetala Mast., 1872 Passiflora mollis HBK., 1817 * nr

and car

900-3000

Uribe 5973 (COL) Prieto 302 (UIS) Idrobo 2037 (COL) Ocampo et al. 54 (HUA) MacDougal 4161 (HUA) Uribe 2568 (COL) Perez-A.669 (COL) Humboldt & Bonpland (P) - Type

LC

Tr V.N.: Granadillita de Monte (ant) Tr

and

2000-3200

LC

and

cun snt

900-1600

VU/EN

Tr Tr Not collected since 1938. Tr Tr V.N.: Granadillo del Magdalena. Tr

and

by cun

1600-2790

EN

and

ant

400-840

NT/VU

and

cl cun to ama ant by cho cq met pu vc ant cl cho qu lg snt to vc

200-1200

NT/VU

amz and

0-710

LC

and

1400-2500

LC/NT

Tr

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Ocampo et al.

Colombian Passifloraceae

-15

Taxon Passiflora monadelpha Jrgensen & HolmNielsen, 1987 nr Passiflora munchiquensis Hernndez (ined), 2003 * nr Passiflora occidentalis Hernndez (ined), 2003 * nr Passiflora panamensis Killip, 1922 Passiflora pilosissima Killip, 1931 * nr Passiflora popayanensis Killip, 1930 * Passiflora punctata L., 1753

Biogeographic Region and

Geopolitical Distribution to vc

Elevation

Collection for Reference Escobar 4859 (HUA)

Bibliographic Reference Holm-Nielsen et al. 1988; Hernndez 2003 Hernndez 2003; [Link] (pers. com.). Hernndez 2003; [Link] (pers. com). Killip 1938

IUCN Category VU/EN Tr

Notes

2800-3310

and

cau vc

1900-3200

Vargas 3909 (HUA)

NT/VU

Tr

and pac

cau cho na pu vc

50-1200

Killip 39025 (COL) Zarucchi et al. 5107 (CHOCO) Lehmann 7630 (US) Lozano 6472 (COL) Romero-C. 3150 (COL)

LC/NT

Tr

pac car

ant bl cho cor

0-500

NT

Tr V.N.: Gulupa (ant) Tr Tr

and and

ant vc cau

1500-2100 2400-2900

Killip 1938 Killip 1938 Croat 1978; Holm-Nielsen et al. 1988

CR VU/EN

and pac

cau cun na vc

20-1750

LC/NT

Tr Tr Collected in the border of Panama and Colombia (cho)

[Passiflora sandrae J. MacDougal, 2006]

pac

cho

800-1100

Garwood 1178 (MO) - Type

MacDougal 2006

Passiflora tribolophylla Harms, 1922 * Passiflora tuberosa Jacq., 1804 nr

pac

ant cau cho

50-1820

Lehmann 5420 (foto, COL) Cuatrecasas 15930 (VALLE) Roldn 2345 (HUA) Plowman 2425 (COL)

Killip 1938; Hno. Daniel 1968 Killip 1938 Killip 1960; Holm-Nielsen et al. 1988; Hernndez 2003 Holm-Nielsen et al. 1988

LC/NT

Tr

and

vc

1200

EN

Tr

Passiflora ursina Killip & Cuatrec., 1960 Passiflora vespertilio L., 1753 Section Hahniopathanthus Passiflora guatemalensis S. Watson, 1887

and

ant na vc

2100-3100

VU/EN

Tr

amz and ori

ama met na

150-500

LC/NT

Tr

and car

ant cl lg ma qu ri to vc

0-1580

Uribe 2532 (COL)

Killip 1938; Ulmer & MacDougal 2004

LC

Tr

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16-

Colombian Passifloraceae

Ocampo et al.

Taxon

Biogeographic Region

Geopolitical Distribution

Elevation

Collection for Reference Killip & Hazen 8670 (Y)

Bibliographic Reference Uribe 1955b; Holm-Nielsen 1974; Ulmer & MacDougal 2004

IUCN Category

Notes Tr V.N.: Granadilla Abroquelada (ant)

Passiflora hahnii (Fourn.) Mast., 1872 nr

and car

ant cl lg ma to vc

100-1250

CR

Section Pseudodysosmia Tr V.N: Pegajosa (qu), Granadilla Culebra (vc), Gulupo (cun). Edible fruit

Passiflora adenopoda Moc. & Sess ex DC., 1828

and car

ant by cl cun cau ma qu ri to vc

100-2100

Cuatrecasas 15703 (VALLE)

Holm-Nielsen et al. 1988; MacDougal 1994

LC

Passiflora lobata (Killip) Hutch. ex J.M. MacDougal,1986

pac

ant cho

0-1200

Gentry 23791 (COL)

MacDougal 1994; Ulmer & MacDougal 2004 Killip 1938; MacDougal 1994.

NT

Tr

Passiflora morifolia Mast., 1872 Section Pseudogranadilla

and

na

500-1000

Karsten s.n. (W) n.v.

EN

Tr

Passiflora bicornis Mill., 1768

car

ant at bl lg ma

0-500

Saravia 3643 (COL)

Killip1938; Holm-Nielsen 1974

LC

Tr V.N.: Cachito de Venado (bl), Cinco Llagas (at)

Passiflora hirtiflora Jrgensen & Holm-Nielsen, 1987 nr Passiflora kalbreyeri Mast., 1883 * Passiflora menispermacea Triana & Planch., 1873 * Section Xerogona

and

ns

2650

Escobar 3152 (HUA) Killip 20284 (COL) Cuatrecasas 9247 (MA)

Hernndez 2003

CR

Tr

and car

ce ns snt

1100-3100

Killip 1938

LC/NT

Tr

and

to

1400-3000

Killip 1938

LC

Tr

Passiflora capsularis L., 1753

and car pac

ant cl cun cho hu lg ma na ns qu snt to vc

100-2000

Uribe 2566 (COL)

Killip 1938; Holm-Nielsen 1974; HolmNielsen et al. 1988 Holm-Nielsen 1974; HolmNielsen et al. 1988

LC

Tr

Passiflora costaricensis Killip, 1922

pac

cho

20-1500

Croat 42591 (HUA)

NT

Tr

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Ocampo et al.

Colombian Passifloraceae

-17

Taxon

Biogeographic Region

Geopolitical Distribution

Elevation

Collection for Reference MacDougal 3823 (HUA) - Isotype Garcia-B. 17279 (COL)

Bibliographic Reference MacDougal 1992; Ulmer & MacDougal 2004 Holm-Nielsen et al. 1988

IUCN Category

Notes

Passiflora escobariana J.M. MacDougal, 1992

and

ant

1090-1100

VU

Tr

Passiflora rubra L., 1753 Subgenus Dysosmia (DC.) Killip, 1938 Passiflora foetida var. eliasii Killip, 1938

and car

ant cl cau cun hu lg pu na pu qu ri to vc

500-2000

LC

Tr V.N.: Chulupa de Monte (cl)

car

at bl ma

0-500

Penell 12029 (N)

Kiliip 1938

VU

Tr V.N.: Flor de la Pasin, Pasionaria (at) Tr V.N.: Granadilla (cho), Flor de la Pasin (at), Gulupo (cun), Bejuco Canastilla (met), Chulupa de Loma (ant hu), Cinco Llagas (cor). I.N.: Ianaleeg murulale (ama). Edible fruit Tr V.N: flor de la pasin (ma), gulupo (cun) Tr V.N.: Flor de la Pasin (vc) Tr V.N.: Flor de la Pasin (ma) Tr Flor de la Pasin (ma)

Passiflora foetida var. gossypiifolia (Desv.) Mast. 1872

amz and car ori pac

ama ant ara at bl by cau ce cor cq cs cun cho gn gv hu lg ma met na ns qu snt suc to va vc

0-1500

Schultes 22576 (COL)

Killip 1938; Martin & Nakasone 1970; Romero-C. 1991; Ulmer & MacDougal 2004; Ulmer & Ulmer, 2005

LC

Passiflora foetida var. hispida (DC.) Killip ex Gleason, 1931 Passiflora foetida var. isthmina Killip, 1938 Passiflora foetida var. moritziana (Planch.) Killip ex Pull, 1937 Passiflora foetida var. sanctae-martae Killip, 1938 * nr Passiflora vestita Killip, 1938 Distephana (Juss.) Killip, 1938

and car

ant bl cun ns to

0-1500

Killip & Smith 21000 (N) Killip 5289 (N) Killip & Smith 21088 (N) Smith 1532 (P) Betancourt 5164 (MO) n.v.

Killip 1938; Ulmer & Ulmer, 2005 Killip 1938

LC

and pac

na snt vc

0-1200

VU

car

ma

0-500

Killip 1938

VU

car

ma

0-500

Kiliip 1938 Killip 1938; Holm-Nielsen et al. 1988

EN

amz

pu

0-500

VU/EN

Tr

Passiflora coccinea Aubl., 1775

amz ori

ama cs gn gv met na va vch

150-1500

Davidse 5321 (COL)

Escobar 1988a

LC

Tr V.N.: Lluvia Padie, Granadillo de Conga (ama), Granadilla colorada (cs). Edible fruit

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18-

Colombian Passifloraceae

Ocampo et al.

Taxon Passiflora involucrata (Mast) A.H. Gentry, 1981 Passiflora glandulosa Cav., 1790 Passiflora quadriglandulosa Rodschied, 1796

Biogeographic Region amz

Geopolitical Distribution ama cq va

Elevation 150-350

Collection for Reference Schultes 6923 (COL) Romero-C. 3668 (AAU) n.v. Lozano 604 (COL)

Bibliographic Reference Escobar 1988a Killip 1938; Holm-Nielsen 1974 Escobar 1988a; Holm-Nielsen et al. 1988

IUCN Category LC Tr

Notes

amz

va

150-500

EN

Tr

amz

ama gu

150-500

LC/NT

Tr Tr V.N.: Granadilla, Oncilla, Parcha de Culebra de Agua (ama) Tr V.N.: Chulupo (cq), Granadilla de Monte (cho), Granadillo (met cq), Gulupa (to). I.N.: Maloca de Fisi (ama). Edible fruit

Passiflora variolata Poepp. & Endl., 1838

amz

ama cq va

150-500

Zarucchi 2197 (COL)

Escobar 1988a

LC/NT

Passiflora vitifolia Kunth, 1817

amz and car ori pac

ama ant bl by cau ce cl cho cor cq cun lg gv ma met na pu ri snt to va vc vch

0-1800

Cuatrecasas 15740 (VALLE)

Killip 1938; Romero C. 1956, 1991; Martin & Nakasone 1970; Holm-Nielsen 1974; HolmNielsen et al. 1988

LC

Subgenus Manicata (Harms) Escobar, 1988 (Syn. Granadillastrum) Passiflora manicata (Juss.) Pers., 1807 Passiflora (Medik.) Mast., 1871 (Syn. Granadilla) Series Digitatae Tr V.N.: Cocorilla (cho). Granadilla, Naracujinha (ama). N.I.: Cipo-Cipo Naracujinha (ama). by cau cl cun na ns qu snt to vc Richter s.n. (COL) Jussieu 1805; Holm-Nielsen 1974; Escobar 1988a Tr V.N.: Tacso (na), Curubo de Monte (qu ns).

and

1400-2700

LC

Passiflora serratodigitata L., 1753

amz and pac

ama cho ns ant

0-1000

Renteria 3542 (COL)

Killip 1938; Holm-Nielsen 1974; HolmNielsen et al. 1988; RomeroC. 1991

LC

Series Laurifoliae Passiflora ambigua Hemsl. ex Hook., 1902 amz and ori pac ant by cl cho cun hu ma met pu snt vc 0-2000 Fuchs 21744 (COL) Holm-Nielsen et al. 1988 LC Tr Edible fruit

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Ocampo et al.

Colombian Passifloraceae

-19

Taxon Passiflora gleasonii Killip, 1924

Biogeographic Region ori

Geopolitical Distribution gn

Elevation 150-500

Collection for Reference Madrin 1014 (COL) Uribe 2405 (COL) Schultes 5875 (US) Zarucchi 1824 (COL)

Bibliographic Reference Killip 1938

IUCN Category EN Tr

Notes

Passiflora guazumaefolia Juss., 1805

and car

ce cor bl ma snt

0-500

Killip 1938; Coppens 2003

LC/NT

Tr V.N: La Parcha (ce), Cocorilla (ma). Edible fruit Tr Tr Edible fruit Tr V.N.: Granadilla (cho met), Granadilla Babosa (na). N.I.: Burucua, Gemarundare, Tuchica, Jino-Goj (va). Edible fruit Tr Reported in the Amazon of Brazil, Peru and Venezuela Tr V.N: Granadilla de Quijos (na), granadilla caucana, curubejo (cau). Cultivated. Edible fruit Tr Edible fruit Tr Edible fruit

Passiflora killipiana Cuatrecasas, 1960 nr Passiflora laurifolia L., 1753

amz and amz pac

cq ama cho cq gv hu met snt va

250-500 0-1700

Killip 1960 Killip 1938

CR LC

Passiflora nitida Kunth, 1817

amz and car ori pac

ama ant cho cq cun cs gn gv ma met na pu va vc

0-1940

Triana 2931 (P)

Killip 1938; Romero-C. 1956, 1991; Holm-Nielsen 1974; Garca-B. 1975; Croat 1978

LC

[Passiflora phellos C. Feuillet, 2004]

amz

90-150

Wurdack & Addeley 43479 (NY) - Holotype

Feuillet 2004

Passiflora popenovii Killip, 1922 nr

and

cau na vc

1200-2050

Escobar & Escobar 1017 (HUA)

Killip 1938; Holm-Nielsen et al. 1988; Romero-C. 1991; Ulmer & MacDougal 2004 Killip 1960 Killip 1938

EW

Passiflora riparia Mart. ex Mast., 1872 Passiflora tolimana Harms, 1894 * Series Incarnatae Passiflora cincinnata Mast., 1868 nr

amz and

cq pu va ant to vc

300-400 820-2000

Smith 3157 (US) Echeverry 3627 (TOLI)

LC/NT NT/VU

and

ns

1200

Killip & Smith 20879 (Y)

Killip 1938

CR

Tr Ornamental (qu). Edible fruit Tr Introduced from Brazil in the 1950s. V.N: Curuba Redonda (ant cl ri qu), Gulupa (cun). Cultivated or feral. Edible fruit

Passiflora edulis f. edulis Sims, 1818

amz and pac

ant cl cau cho cun gv met na qu ri snt to vch vc

1100-2750

Idrobo 1637 (COL)

Holm-Nielsen et al. 1988; Vanderplank 2000; Ulmer & MacDougal 2004

NE

Biota Colombiana 8 (1), 2007

20-

Colombian Passifloraceae

Ocampo et al.

Taxon

Biogeographic Region

Geopolitical Distribution ant ara bl ce cl cho cun gn hu met pu qu snt ri to vc

Elevation

Collection for Reference

Bibliographic Reference Killip 1938; Ulmer & MacDougal 2004

IUCN Category

Notes Tr Introduced from Brazil in the 50s. V.N.: Maracuy. Cultivated. Edible fruit

Passiflora edulis f. flavicarpa Degener, 1932 nr

amz and car ori pac

0-1800

Silvestone 14399 (CUVC)

NE

Series Kermesinae Passiflora lehmanni Mast., 1885 * amz and ant cau cl cun hu pu qu ri snt vc cun ma qu snt to vc 1000-2000 Uribe 2588 (COL) Killip & Smithii 15015 (MO) - Holotype Marulanda 91 (HUA) Killip 1938; Holm-Nielsen 1974 Killip 1938; Holm-Nielsen 1974 Killip 1938; Hno. Daniel 1968 LC Tr

Passiflora smithii Killip, 1930

and car

500-2000

LC

Tr V.N.: Curuba Silvestre (to).

Passiflora trisulca Mast., 1887 * Series Lobatae

and

ant cl vc

1300-1800

NT

Tr

Passiflora caerulea L., 1753 nr

and

cl cun qu

1000-2700

Ocampo 83 (VALLE)

Deginani 2001

Tr Introduced from Argentina. Ornamental. Edible fruit LC/NT Tr Tr Collected on the border of Ecuador and Colombia (na) DD NT/VU Tr Tr

Passiflora gritensis H. Karst., 1859 [Passiflora montana Holm-Nielsen & Lawesson, 1987] Passiflora picturata Ker, 1822 nr Passiflora pennellii Killip, 1924 *

and

by ns

2450-2500

Cuatrecasas 1808 (COL) Holm-Nielsen et al. 6200 (AAU) Uribe 1334 (US) Uribe 4827 (COL) Marulanda & Mrquez 1665 (HUA)

Killip 1938

and

2600

Holm-Nielsen et al. 1988

ori and

met ant cun

450 1200-1600

Killip 1938, 1960 Killip 1938 Killip 1938; Holm-Nielsen 1974;HolmNielsen et al. 1988 Killip 1938 Killip 1938; Holm-Nielsen 1974

Passiflora resticulata Mast. & Andr, 1884

amz and pac

cau gv na vc

0-2000

NT/VU

Tr

Passiflora semiciliosa Planch & Linden, 1873 * Passiflora subpeltata Ortega, 1798

and car

ma ns ant bl cau ce cor cq cs cun ma suc to vc

1850-3000

Garcia-B. 20749 (COL) Ramrez et al. 11507 (CAUP)

VU

Tr V.N.: Gulupa, Palcha (ns) Tr V.N.: Cocorilla (bl)

amz and car ori

0-2400

LC

Biota Colombiana 8 (1), 2007

Ocampo et al.

Colombian Passifloraceae

-21

Taxon Series Quadrangulares

Biogeographic Region

Geopolitical Distribution

Elevation

Collection for Reference

Bibliographic Reference

IUCN Category

Notes

Passiflora alata Curtis, 1788 nr

amz

ama

200

Ocampo 82 (VALLE)

Killip 1938; Ulmer & MacDougal 2004

DD

Tr Introduced (qu vc) from Brazil in the 90s. V.N.: Maraca. Cutivated. Edible fruit Tr V.N.: Badea (ant cl hu cun met qu ri), corvejo (snt), Granadillo Grande (cau), curuba (vc), Motorro (gn). Cultivated. Edible fruit

Passiflora quadrangularis L.,1759

amz and car ori pac

ama ant bl cau cl cho cq cun gn hu ma met na ns qu va ri snt to vc

0-1500

Gentry 15371 (COL)

Killip 1938; Romero-C. 1956, 1991; HolmNielsen1974; Holm-Nielsen et al. 1988

LC

Series Menispermifoliae Gerlach & Ulmer, 2000; Ulmer & MacDougal 2004 Croat 1978; Holm-Nielsen et al. 1988

Passiflora chocoensis G. Gerlach & T. Ulmer, 2000 * nr

pac

cho

0-100

Gerlach 434917 (COL) - Holotype

CR

Tr

Passiflora menispermifolia Kunth, 1817 Series Simplicifoliae

amz car and pac

ant bl by cho cor cq cun met na ns snt to vc

0-2140

Cuatrecasas 15541 (VALLE)

LC

Tr V.N.: Chulupe (cq)

Passiflora danielii Killip, 1960 *

and

ant

1300-2600

Hno. Daniel 1536 (MEDEL) - Isotype Sanchez 17 (COL) Cadena 83 (UIS)

Killip 1960; Hno. Daniel 1968

VU/EN

Tr

Passiflora longipes Juss., 1805 * Passiflora longipes var. oxyphylla L. Uribe, 1977 *

and

cun by qu snt to

2500-3500

Killip 1938

NT

Tr

and

by ns snt

2000-2600

Uribe 1977

NT

Tr

Passiflora oerstedii Mast., 1872

and ori pac

ant cau cho cun cau met na qu ri vc

0-2000

Romero-C. 6141 (COL)

Killip 1938; Holm-Nielsen 1974; HolmNielsen et al. 1988

LC

Tr

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Ocampo et al.

Taxon Series Tiliaefoliae Passiflora ligularis f. lobata (Mast.) Killip, 1938 nr

Biogeographic Region

Geopolitical Distribution

Elevation

Collection for Reference

Bibliographic Reference

IUCN Category

Notes

and

ant

1800-2000

Archer 1498 (COL)

Killip 1938

NT

Tr V.N: Granadilla. Cultivated. Edible fruit Tr V.N: Granadilla; Granadilla Pipo (na). N.I.: Awapit (na). Cultivated. Edible fruit

Passiflora ligularis Juss., 1805

and

ant cl cun cau by cho hu met na ns pu qu ri snt to vc

1550-2500

Dombey 739 (P) - Type

Killip 1938; Romero-C. 1956, 1991; Holm-Nielsen et al. 1988

LC

Passiflora magnifica L.K. Escobar, 1990 *

and

ant

1250-1750

Callejas 6586 (HUA) n.v.

Escobar 1990; Ulmer & MacDougal 2004

VU

Tr Edible fruit

Passiflora maliformis L., 1753

and car pac

ant by cl cau cun cho hu ma na qu snt to vc

0-2200

Humboldt & Bonpland 1804 (P) - Type

Killip 1938; Romero-C, 1956, 1991; Holm-Nielsen, 1974; Garca-B. 1975; HolmNielsen et al. 1988

LC

Tr V.N: Gulupa, Granadilla de Piedra, o de Hueso (cu, na vc), Gurapa (snt), Chulupa (hu). Cultivated. Edible fruit Tr V.N: Palchita (ns). Edible fruit Tr V.N.: Granadilla (cho), Camelo (vc). Cultivated. Edible fruit

Passiflora multiformis Jacq., 1809

and car

lg ma ns

0-1300

Romero-C. 8992 (COL)

Killip 1938

NT/VU

Passiflora palenquensis Holm-Niels. & Lawesson, 1987

pac

ant cau cho na vc

0-1200

Espina & Garcia 1951 (COL)

Holm-Nielsen & Lawesson 1987; HolmNielsen et al. 1988

LC

Passiflora platyloba Killip, 1922

pac

cho

0-1050

Gentry & Juncosa 40946 (COL)

Gentry 1976

NT/VU

Tr Edible fruit

Passiflora seemannii Griseb., 1858

amz and pac ori

ant by cho cun gn gv met ns snt va vc vch

0-1300

MacDougal 4144 (HUA)

Croat 1978

LC

Tr V.N.: Palcha, Chulupa (met), Granadilla Montaera (cun). Edible fruit

Biota Colombiana 8 (1), 2007

Ocampo et al.

Colombian Passifloraceae

-23

Taxon

Biogeographic Region

Geopolitical Distribution

Elevation

Collection for Reference

Bibliographic Reference

IUCN Category

Notes Tr V.N.: Guayabita Cimarrona (ma). Edible fruit Tr V.N.: Granadilla, Machimbi (Colombia). Cultivada. Fruto comestible.

Passiflora serrulata Jacq., 1767

car

at ma lg

0-500

Bunch 601 (FMB)

Killip 1938; Coppens 2003

NT/VU

Passiflora tiliifolia L., 1753

and pac

ant cau cho cl na qu vc to

1100-2500

Gonzlez 1411 (CAUP)

Killip 1938; Holm-Nielsen 1974; HolmNielsen et al.1988; Coppens 2003

LC/NT

Subgenus Porphyropathanthus L.K Escobar, 1989 Passiflora sierrae L.K. Escobar, 1989 * Subgenus Psilanthus (DC.) Killip, 1938 Passiflora bicuspidata (H. Karst.) Mast., 1872 * Passiflora hyacinthiflora Planch. & Linden, 1873 * Passiflora trinervia (Juss.) Poir., 1811 * Subgenus Rathea (Karst.) Killip, 1938 Passiflora andina Killip, 1938 nr Killip 1938; Holm-Nielsen et al.1988 Escobar 1986, 1988 and by cun ns snt 2500-3500 Rojas 138 (CDMB) Garcia-B. 20700 (COL) Cuatrecasas 20241 (VALLE) Uribe 1972; Killip 1978 Killip 1938 VU Tr car ma 3000-3700 Cuatrecasas 24375 (COL) Escobar 1989 EN/CR Tr

and

by ma ns

2900-3300

LC/NT

Tr

and

cl qu to vc

2500-3700

Jussieu 1805; Killip 1938

VU

Tr

and

na

2800

Karsten (V)

CR

Tr

Passiflora colombiana L.K. Escobar, 1986 * Subgenus Tacsonia (Juss.) Tr. & Planch, 1873 Section Bracteogama Passiflora cumbalensis var. caucana L.K. Escobar, 1987 *

and

na pu

3000-3600

Mora 6175 (PSO) Paratype

CR

Tr

and

cau

2300-2800

Tryon 6001 (COL)

Escobar 1987, 1988b

LC

Tr V.N.: Curuba de Monte. Edible fruit

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Taxon

Biogeographic Region

Geopolitical Distribution

Elevation

Collection for Reference

Bibliographic Reference Romero-C. 1956; HolmNielsen 1974; Escobar 1987, 1988; HolmNielsen et al. 1988

IUCN Category

Notes

Passiflora cumbalensis var. cumbalensis (H. Karst.) Harms,1894

and

na pu

3000-3800

Fernandez 5834 (COL)

LC/NT

Tr V.N.: Curuba Roja, Tauso (na). Edible fruit

Passiflora cumbalensis var. goudotiana (Triana & Planch.) L.K. Escobar, 1987

and car

ant by cl cq cun hu ma na pu qu ri snt to vc

1800-3300

Uribe 2593 (COL)

Escobar 1987, 1988; HolmNielsen et al. 1988

LC

Tr V.N.: Curuba bogotana (cun), Curubo mucura, curuba rosada, Tausa (na). Cultivated. Edible fruit Tr Reported in the northern Andes of Ecuador Tr V.N.: Curuba. Edible fruit (by cun) Tr V.N.: Curuba de Castilla (ant by cu cl); Tauxso (na). Cultivated. Edible fruit Tr V.N.: Curuba India. Cultivated. Edible fruit.

[Passiflora sanctae-barbarae Holm-Nielsen & Jrgensen, 1987] Passiflora tripartita var. azuayensis Holm-Nielsen & Jrgensen, 1988 nr

and

2200-2700

Harling & Andersson 12445 (AAU) - Isotype Escobar 19999 (HUA)

Holm-Nielsen et al. 1988

and car

ant by cun ma ns

2000-2610

Holm-Nielsen et al. 1988

LC/NT

Passiflora tripartita var. mollissima Holm-Nielsen & Jrgensen, 1988

and car

ant by cau cl cun ma na ns pu snt vc

2200-3500

Romero-C 8007 (PSO)

Holm-Nielsen et al. 1988; Romero-C. 1991; Ulmer & MacDougal 2004

LC

Passiflora tarminiana Coppens & Barney, 2001

and

ant by cau cl cun hu na qu snt to vc

2000-2900

Coppens 72 (COL) - Type

Coppens et al. 2001; Campos 2001

LC

Section Colombiana Series Colombianae Passiflora adulterina L.f., 1781 * and by cun snt to 2600-3600 Barclay 4517 (COL) Escobar 1988a NT Tr Tr V.N.: Curuba Paramera (cun)

Passiflora crispolanata [Link], 1954 *

and

by cun

2500-3500

Uribe 6773 (COL)

Uribe 1954; Escobar 1988a

NT

Passiflora cuatrecasasii Killip, 1960 *

and

by cun met snt

2200-3500

Cuatrecasas 9479 (foto, MEDEL)

Killip 1960; Escobar 1988a

VU

Tr

Biota Colombiana 8 (1), 2007

Ocampo et al.

Colombian Passifloraceae

-25

Taxon Passiflora formosa T. Ulmer, 1999 *

Biogeographic Region and

Geopolitical Distribution by

Elevation 3000-3100

Collection for Reference Uribe 5945 (COL)

Bibliographic Reference Ulmer 1999 Jussieu 1805; Holm-Nielsen 1974; Escobar 1988a

IUCN Category EN Tr

Notes

Passiflora lanata (Juss.) Poir., 1811 *

and

cun by snt to

2200-3500

Uribe 2587 (COL)

NT/VU

Tr V.N.: Granadilla (cun)

Passiflora pamplonensis Planch.& Linden ex Triana & Planch., 1873 * Passiflora rigidifolia Killip, 1960 * nr Passiflora rugosa var. rugosa (Mast.) Triana & Planch., 1873 Passiflora rugosa var. venezolana L.K. Escobar, 1986 Passiflora trianae Killip, 1938 * Passiflora truxillensis Planch. & Linden, 1873 nr Series Leptomischae

and

snt

2000-3000

Funck & Schlim 1385 (foto, VALLE) Burke 185 (K) - Type

Escobar 1988a

EN/CR

Tr Curubita de Piuela (snt) Tr Known only from the type.

and

ant

3750

Killip 1960

DD

and

cun met ns

3000-3500

Peuela 008 (COL)

Escobar 1988a

LC/NT

Tr

and

ns snt

2500-3500

Garcia-B. 20001 (COL) Escobar 569 (COL) V. Barney & G. Coppens (foto), pers. com.

Escobar 1988a

LC/NT

Tr

and

ns snt

3000-3500

Escobar 1988a

VU/EN

Tr

and

ns

1800-3000

Escobar 1988a; Ulmer & Ulmer 2005

EN

Tr

Passiflora antioquiensis H. Karst., 1859 *

and

ant cau cl cun hu pu qu ri to vc

1800-2700

Escobar 2133 (HUA)

Hno. Daniel 1968; Uribe 1972; GarciaB. 1975; Escobar 1988a

LC/NT

Tr V.N.: Granadilla (vc), Curuba Antioquea (ant). Wild or cultivated in home gardens. Edible fruit Tr Known only from the type. Tr V.N.: Curuba de Monte (cl qu ri). Edible fruit Tr Edible fruit

Passiflora cremastantha Harms, 1922 *

and

cau

2000-2500

Lehmann 5421 (F) - Type

Escobar 1988a

EX

Passiflora flexipes Triana & Planch., 1873 *

and

cl qu ri

2500-3380

Vargas 626 (FAUC)

Escobar 1988a

NT/VU

Passiflora leptomischa Harms, 1922 *

and

ant cau qu vc

2000-2800

Escobar et al. 4421 (PSO)

Escobar 1988a

LC/NT

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26-

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Taxon

Biogeographic Region

Geopolitical Distribution

Elevation

Collection for Reference Escobar 4853 (COL)

Bibliographic Reference Escobar 1988a, 1989b; Campos 2001

IUCN Category

Notes Tr V.N.: Curuba de Monte (vc). Edible fruit

Passiflora tenerifensis L.K. Escobar, 1988 * Series Quindiensae Passiflora linearistipula L.K. Escobar, 1988 * Passiflora quindiensis Killip, 1938 * Section Fimbriatistipula Passiflora fimbriatistipula Harms, 1894 * Passiflora uribei L.K. Escobar, 1988 * Section Parritana Passiflora jardinensis L.K. Escobar, 1988 * Passiflora parritae (Mast.) L.H. Bailey, 1916 * Section Poggendorffia

and

vc

2800-3100

EN/CR

and

cl

2650-3170

Ocampo et al. 56 (HUA) Uribe 3320 (COL)

Escobar 1988a Escobar 1988; Campos 2001

EN/CR

Tr Not collected since 1984. Tr

and

to

2900-3100

VU/EN

and

cau hu

2130-3240

Fernandez et al. 30182 (AFP) Escobar et al. 2896 (HUA)

Escobar 1988a Uribe 1958; Escobar 1988a

NT/VU

Tr I.N.: Pachuaca (cau) Tr

and

na pu

2500-2700

EN

and

ant

2750-3000

Zarucchi 6963 (COL) Snchez 15 (FAUC)

Escobar 1988b

VU/EN

Tr Tr Curuba de Monte (to). Edible fruit

and

cl qu ri to

2500-3020

Escobar 1988a

VU/EN

Passiflora pinnatistipula Cav., 1799

and

ant by cun na ns

2000-3600

Uribe 6643 (COL)

Escobar 1988b; Holm-Nielsen 1974; HolmNielsen et al. 1988; Campos 2001

LC/NT

Tr V.N.: Curuba Redonda, Gulupa (cun) Cultivated. Edible fruit Tr Natural hybrid of P. pinnatistipula x P. tripartita var. mollissima. Edible fruit (when fertile)

Passiflora x rosea (H. Karst.) Killip, 1938

and

by cun

2500-3500

Uribe 3941 (COL)

Escobar 1988a

VU

Section Tacsonia Tr V.N.: Curuba de Monte (vc), Curubo de Pramo (cun), Palchuaca (cau), Curubito de Indio (cl). Edible fruit

Passiflora mixta L. f., 1781

and

ant cl by cau cun na ns qu ri snt to vc

1700-3700

Humboldt & Bonpland (P) - Type

Escobar 1988a; Holm-Nielsen 1974; HolmNielsen et al. 1988; Garca-B. 1975

LC

Biota Colombiana 8 (1), 2007

Ocampo et al.

Colombian Passifloraceae

-27

Taxon

Biogeographic Region

Geopolitical Distribution

Elevation

Collection for Reference

Bibliographic Reference Holm-Nielsen 1974; Escobar 1988a; RomeroC. 1991; Coppens 2003

IUCN Category

Notes

Passiflora schlimiana Triana & Planch., 1873 *

car

ce lg ma

2400-3220

Romero-C. 7407 (COL)

VU/EN

Tr V.N.: Curuba. Edible fruit

Section Tacsoniopsis Passiflora bracteosa Planch. & Linden, 1873 Passiflora purdiei Killip, 1938 * Subgenus Tryphostemmatoides (Harms) Killip), 1938 Passiflora tryphostemmatoides Harms, 1894 Passiflora gracillima Killip, 1924 Passiflora arbelaezii L. Uribe, 1957 Passiflora pacifica L.K. Escobar, 1988 * and ant cau hu qu ri vc ant cau cl hu na qu to ant cau cho cun na vc cho na vc 1000-2700 Lehmann 5662 (K) - Isotype Penell 9393 (MO) - Isotype Roldn 1162 (COL) Escobar 2143 (HUA) Killip 1938; Holm-Nielsen et al. 1988 Killip 1924, 1938 NT Tr and ns snt 2200-3000 Garcia-B. 20745 (COL) Purdie s.n. (K) n.v. Escobar 1988a EN Tr V.N.: Palchoaca (ns, snt) Tr Known only from the type.

and

cun ma

Escobar 1988a

EX

and

2000-3150

LC

Tr V.N. Tr V.N.: Golondrina (cho) Tr

and pac

0-2300

Uribe 1957

LC/NT

pac

0-1800

Escobar 1988b

LC/NT

Discussion Colombia has been subject of many studies focused on inventories of plant species groups (Gentry 1993; Silverstone-Sopkin & Ramos 1995; Galeano et al. 1998; Rangel 1995, 2002). Passifloraceae have been inventoried in taxonomical works by Escobar (1998a, 1989, 1990 inedited) and Hernndez & Bernal (2000). Compared to the latter, we have added new information on geographical distribution of each taxon and extended the list to a total of 167 Passifloraceae species, from three genera and the five biogeographic regions, with reports of 26 species new to Colombia. For obvious reasons, the quality of botanical inventories depends on the quality of taxonomical work in this complex family. While the definition of genera and subgenera should not significantly affect studies of the distribution of its diversity across the Colombian territory, such work may be affected to some extent by poor definitions below the subgenus level. Indeed, several morphological groups include species that are very si-

milar, and regularly reported as very difficult to distinguish from each other. In several cases, experts may have underestimated intraspecific variation in widely distributed species, or even intra-individual variation, splitting well-known species in several new species only distinguished by a few quantitative or color traits. Among the difficult groups, let us mention particularly subgenus Astrophea, whose species tend to be less well differentiated, at least in sterile specimens, by the position and number of the nectar glands, having only two at the junction of the lamina and petiole, while they may show impressive intraspecific variation in pubescence and intra-individual variation in leaf size and shape according to light exposure and whole tree development (heteroblasty). Also in the subgenus Decaloba there are several morphological groups that demand great experience and care in their identification, even for the most common species such as P. capsularis and P. rubra, which can be found in the same habitats. In the most difficult cases, several species have even changed status several times. For instance, Killip
Biota Colombiana 8 (1), 2007

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merged P. bauhinifolia Kunth. with P. andreana Mast. in 1938, and restored it as a distinct species in 1960, while Holm-Nielsen et al. (1988) merged P. bauhinifolia with another close relative, P. alnifolia, a position we have adopted here. A couple of other species, such as P. mollis and P. cuspidifolia or P. hahnii and P. guatemalensis, may also show very little morphological difference, but differ in their altitudinal distribution, which confirms they are different. Many new species of subgenus Distephana are also questionable, as one of its two most common species, P. coccinea Aubl., distributed in most of the Amazon, has been split in several species on the basis of bract size, number of nectar glands, and small variation in numbers and respective colors of the corona series. Concerning Colombia, Vanderplank (2006) underlined that the description of P. coccinea by Escobar (1988) matches perfectly that of P. miniata Vanderplank, so he considered the latter a Colombian species. However, we have not adhered to this opinion for several reasons: Vanderplank described it on material grown in glasshouse and his report does not refer to the examination of Colombian materials. The type and level of the differentiation described between the various new species and P. coccinea is at most of the same order as morphological variation in other common widespread species (e.g. P. vitifolia, P. foetida, P. suberosa, P. alnifolia, P. capsularis, P. mixta, P. cumbalensis, P. maliformis, or P. emarginata). He reported a high level of sexual compatibility with the other common Distephana species, P. vitifolia, which raises the expectation of sexual compatibility with the even closer true P. coccinea. Thus we have stuck to the treatment of P. coccinea by Escobar (1988), whose quantitative description is more precise than the original by Aublet (1775), but not fundamentally different. Within subgenus Passiflora, P. maliformis, P. serrulata and P. multiformis constitute other cases of possible overclassification, as they are mostly differentiated by the degree of lobation of their leaves, a trait that is quite variable in many other species, including other Tiliifoliae, such as P. ligularis (Killip 1938; pers. obs.). A wider problematic group is the series Laurifoliae, with ten species in Colombia, always difficult to identify from incomplete specimens. Although they probably constitute a very young group and they exhibit a high number of common traits, species of subgenus Tacsonia are relatively easy to differentiate. Particularly interesting are the endemics of Colombian section, from the center of the cordilleras, often characterized by a very long peduncle and linear-lanceolate stipules, and from the northeast and up to the Venezuelan Andes. Several authors have reported easy interspecific hybridization in subgenus Tacsonia, involving cultivated, as well as wild materials (Escobar 1985). This phenomenon,
Biota Colombiana 8 (1), 2007

by producing spontaneous off-types, may have led to some overclassification in this subgenus. Indeed, of the 30 species reported here for Colombia, five are known only from the type material (P. cremastantha Harms, P. formosa Ulmer, P. pamplonensis Planch.& Linden ex Triana & Planch., P. purdiei Killip, P. rigidifolia Killip) are known only from the type material. Whether this is due to high endemism, ancient extinction, or off-types resulting from hybridization cannot be ascertained, unless a second specimen is recorded, as we did for P. linearistipula. It is important to note that P. formosa was described as a new species from the same specimen considered an off-type of P. lanata (Juss.) by Escobar (1988). Overclassification may be suspected even in better known species, as P. parritae (Mast.) Bailey, and P. jardinensis L.K. Escobar. Indeed, in populations of the former, we have observed sufficient morphological variation to include the few known specimens of the latter species, which might simply represent a small isolated population. On the other hand, most endemics of subgenus Tacsonia were found in difficult to access highlands, and more species can be described from relatively poorly explored areas such as the South of Tolima, Santander and Norte de Santander departments. Our list ranks Colombia as the country with the highest richness of Passifloraceae, followed by Brazil with 127 species. Figure 4 allows comparisons for species richness and relative diversity of passion flowers in the Neotropics, showing the strong influence of latitude (typical of a tropical distribution) and topography on Passiflora diversity. Colombian species richness and diversity is more than twice that of Peru and Venezuela, two countries of similar surface and latitude. Given its much smaller area, Ecuador also presents an impressive diversity. Thus, the northern Andes of Colombia and Ecuador clearly constitute the center of diversity for the genus Passiflora. This is probably due to the greater availability of habitats, especially at high elevations, in these two countries. The presence of three Andean cordilleras in Colombia very probably played a significant role. Indeed, radiation has been very active in the northern Andes, with particular contribution of recent and fast evolving groups, such as subgenera Rathea and Tacsonia, accounting for more than 41 highland species in Colombia and Ecuador. Among them, 21 (14%) species are endemic to Colombia. Colombian highlands are also rich in representatives of subgenus Decaloba. According to Escobar (1988a), 40% of the New World Passifloraceae are found in the Andes. In Colombia, habitats between 1000 and 3000 m account for only 27% of the land area, yet 81% of the species of Passifloraceae grow there. With 123 species, the Andean region concen-

Ocampo et al.

Colombian Passifloraceae

-29

Figure 4. Distribution of species richness of Passifloraceae in American countries, according to information gathered from Killip (1938, 1960), Escobar (1988, 1989, 1990 inedited, 1994), Holm-Nielsen et al. (1988), Jrgensen & Len (1999), MacDougal (1994), Vanderplank (2000), Deginiani (2001), Tillet (2003), Ulmer & MacDougal (2004), records of the herbaria cited in this study and many journal articles related with the description of new species present in the America.

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trates the highest richness, mainly between 1000 and 2000 m. The Caribbean region shares the highest proportion of species (27) with the Andean region (Box 1). This is mostly due to the presence of the Sierra Nevada de Santa Marta mountain range in northern Colombia, with a steep gradient of elevation from the Caribbean Sea to 5775m summits. The increase of species richness and endemism with the elevation is generally interpreted as a result of the increasing isolation and decreasing habitat surface in high mountain regions, leading to small, fragmented populations which are prone to speciation (Simpson 1975; Jrgensen et al. 1995). Another contribution to the particular species richness in Colombia and Ecuador is that of the Pacific Coast region, which continues down from the similar highly diverse ecosystems of Central America (Choc-Darin/Western Ecuador hotspot of Myers et al. 2000). In strong contrast with the conditions prevailing in the westerns Andes and the Peruvian coast that are arid or semi-arid, or the drier and more contrasted climate of Venezuela, this area receives one of the highest precipitation rates in the world. The composition of the Passifloraceae species of this region appears both diverse and well-differentiated when compared to that of the other biogeographic regions (Figure 2). This is not surprising, considering that the Choco region is recognized as one of the most diverse biotas in the world, with nearly 40% endemism (Gentry 1986). Until recently, the genera Dilkea and Ancistrothyrsus were only known as originating from the Amazon basin; however, Escobars description of A. antioquiensis (1990 ined.) in the Andes and the observation of Dilkea retusa in the Andes and Pacific regions extend their distribution to other important biota. The distribution of Passifloraceae has been drastically affected by deforestation, principally in the Andean region. Its historical range corresponds to a region with a long history of livestock and agriculture that now supports extensive coffee, sugar cane, rice, banana, and potato plantations. According to our field observations, very common species, such as P. adenopoda, P. alnifolia, P. capsularis, P. coriaceae, P. rubra, P. suberosa, and P. mixta, are mostly species that thrive in secondary forests or disturbed old-growth forests. Human disturbances may even have contributed to the extention of their distribution, as reported with other plants (Svenning 1998). According to Myers et al. (2000) and Robbirt et al. (2006), rarity and endemism represent two factors of particular significance in the consideration of the risk
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of decline and extinction. In this context, most Colombian Passifloraceae (70.6%) are under some degree of threat according to IUCN criteria. Only 29.4% (48 species) fall in the least concern category (LC), which clearly illustrates the alarming situation for the family (Figure 3). Our results are consistent with a first Red List of Colombian Plants published by the von Humboldt Institute (Caldern 2005), based on the 141 species listed by Hernndez & Bernal (2000), with similar percentages for each category. However, this list only includes P. colombiana L.K Escobar under the category of critically endangered species (CR), while ours places 16 species in this category. A second list, recently published by Hernndez & Garca (2006), includes two different species, P. cremastantha and P. pamplonensis, in this category. Despite several attempts by Escobar and ourselves, the former species, collected before 1922, is only known from the type specimen. Escobar (1988) was followed in considering its probable extinction. Moreover, the list of Hernndez & Garca (2006) gives much lower numbers for the other threat categories, placing as few as 25 species in the threat categories (including two species in the NT category) and 119 species in the Least Concern one. These numbers are far from likely for a group which (i) exhibits its highest diversity in the highly disturbed central coffee growing zone and (ii) includes 58 endemics. The general discrepancy is probably due the fact that our extensive inventory and direct field observations allowed us to take into account both the number of records and existing populations, as well as the date of the last record for each species, evidencing their dramatic reduction over the recent period. Exploration for Passifloraceae was not possible in the protected areas of Colombia that are of essential importance for the conservation of the countrys biodiversity, as the Colombian Ministry of the Environment (MMA) denied us permission to access. Another limiting factor of research for conservation purposes is the armed conflict prevailing in many parts of the country (Martin & Szuter 1999; Dvalos 2001). Forests in the northern Andes are currently one of the major conservation priorities on a global scale due to their fragility, biological richness, high rates of endemism and multiple anthropogenic threats (Olson & Dinerstein 1998). As Passifloraceae display very high species richness, endemism and risk of extinction in this area; and given their multiple ecological interactions with many organisms, as well as their economic potential, this family should constitute both an important conservation target, as well as a good indicator of the success of the efforts made.

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Conclusions With 167 reported species, Colombia is the country with the highest Passifloraceae richness. This richness is concentrated in the Andean region, particularly in the departments of Antioquia, Valle del Cauca and Cundinamarca. Comparing data with other countries confirms that the northern Andes of Colombia and Ecuador constitute the center of diversity for the most important genus, Passiflora. The limited number of explorations in parts of the Andes, the Amazonian and the Orinoquean regions raises expectations that Colombia may harbor many, as yet, unknown species. Future

studies should encompass new regions, including protected areas and areas of conflict. Indeed, more information about the species diversity and its distribution is urgently required for the in situ conservation of, both, species and habitat. Both aspects may even be combined if the genus Passiflora can be used as an indicator of biodiversity in the Andean region, as was the objective of a project in the coffee growing area. Another important aspect is its direct valorization as a germplasm resource for crop diversification programs, implying the need for a better understanding of its morphological and genetic diversity.

Acknowledgements
The authors wish to thank the herbaria that provided specimens or collection data, particularly Francisco J. Roldn (HUA) and Alexandra Hernndez (COL), as well as Colciencias, the Colombian Ministry of the Environment (MMA) and the Research Center of the Colombian Coffee Grower Federation (Cenicaf) for funding the collection missions. The first author gratefully acknowledges financial support from the Gines-Mera Fellowship Foundation (CIAT-CBN). We are indebted also to Jos O. Velasquez (Casa Mutis, Mariquita), Hernando Criollo ([Link]), Mauricio Villegas (Cenicaf), Vicky Barney (Bioversity International), Alvinxon Castro ([Link]), Robinson Galindo (PNN Catatumbo), Carolina Alcazar (Proselva) and Gustavo Morales (J.B. Jos Celestino Mutis) for assistance in obtaining plant data for this study. We are extremely grateful to Colombian farmers contacted in the fieldwork for their continuous help and availability in localizing a great part of the observed plant material.

Literature cited
Apple J., D. Feener. (2001). Ant visitation of extrafloral nectaries of Passiflora: the effects of nectary attributes and ant behavior on patterns in facultative ant-plant mutualisms. Oecologia 127(3): 409-416. Aublet F. (1775). Histoire des plantes de la Guiane franaise. Paris : P.F. Didot, 2: 828-324. Brooks T. M., R.A. Mittermier., C.G. Mittermier., G.A.B. da Fonseca., A.B. Rylands., W.R. Konstant., P. Flick., J. Pilgrim., S. Oldfield., G. Magin., C. Hilton-Tyalor. (2002). Habitat loss and extinction in hotspot of biodiversity. Conservation Biology 16: 909-923. Bchert A., J. Mogens. (2001). The fragility of extreme specialization: Passiflora mixta and its pollinating hummingbird Ensifera ensifera. Journal of Tropical Ecology 17: 323-329. Cavanilles A.J. (1790). Monadelphia classis dissertation decen. Diss. 10: Decemia dissertatio botnica de Passiflora. Typographia regia, Madrid: 439-463. Caldern E. (2005). Familia Passifloraceae. Programa de biologa de la conservacin, proyecto flora amenazada. Instituto Alexander von Humboldt. [Link] Cited 3 abril de 2005. Campos T. (2001). La Curuba: Su cultivo. Bogot, Colombia, IICA. 30pp. Coppens dEeckenbrugge G., V. Barney, P.M. Jrgensen, J.M. MacDougal. (2001). Passiflora tarminiana, a new cultivated species of Passiflora subgenus Tacsonia (Passifloraceae). Novon 11: 8-15. Coppens dEeckenbrugge G. (2003). Promesas de las pasifloras. Memorias del X Seminario Nacional y IV Internacional sobre Especies Promisorias, Medelln Octubre 29-31 de 2003. CD. Croat T. (1978). Flora of Barro Colorado Island. Stanford University Press, Stanford. 943pp. De Candolle A.P. (1828). Mmoires de la Socit de Physique et dHistoire Naturelle de Geneve 1 : 434-436. Deginani N.B. (2001). Las especies argentinas del gnero Passiflora (Passifloraceae). Darwiniana 39: 43-129. Dvalos L.M. (2001). The San Lucas mountain range in Colombia: how much conservation is owed to
Biota Colombiana 8 (1), 2007

32-

Colombian Passifloraceae the violence?. Biodiversity and Conservation 10: 69-78.

Ocampo et al.

Dharwan K., S. Dharwan, A. Sharma. (2004). Passiflora: a review update. Journal Ethno-Pharmacology 94: 1-23. Endress P.K. (1994). Diversity and evolutionary biology of tropical flowers. Cambridge, England: Cambridge University Press. Escobar L.K. (1985). Biologa reproductive de Passiflora manicata e hibridacin con la curuba Passiflora mollissima (H.B.K.) Bailey. Actualidades Biolgicas. 14 (54): 111-121. Escobar L.K. (1986). New species and varieties of Passiflora (Passifloraceae) from the Andes of South America. Systematic Botany 11(1): 88-97. Escobar L.K. (1987). A taxonomic revision of the varieties of Passiflora cumbalensis (Passifloraceae). Systematic Botany 12 (2): 238-250. Escobar L.K. (1988a). Passifloraceae. Flora de Colombia 10. Instituto de Ciencias Naturales, Universidad Nacional de Colombia, 138pp. Escobar L.K. (1988b). Novedades en Passiflora (Passifloraceae) de Colombia Mutisia 71:1-8. Escobar L.K. (1989). A new subgenus and five new species in Passiflora (Passifloraceae) from South America. Annals of the Missouri Botanical Garden 76: 877-885. Escobar L.K. (1990). Two new species of Passiflora (Passifloraceae) from northern South America. Phytologia 69 (5): 364-365. Escobar L.K. (1990, indito). Una revisin taxonmica de Passiflora subg. Astrophea (Passifloraceae). 289 pp. Escobar L.K. (1994). Two new species and a key to Passiflora subg. Astrophea. Systematic Botany 19 (2): 203-210. Feuillet C. (2004). Passiflora phellos, a new species in subgenus Passiflora (Passifloraceae). Novon 14: 285-287. Feuillet C., J.M. MacDougal. (2002). Checklist of recognized species names of passion flowers. Passiflora 12 (2) : 41-43. Feuillet C., J.M. MacDougal. (2003). A new infrageneric classification of Passiflora L. (Passifloraceae). Passiflora 13 (2): 34-38.
Biota Colombiana 8 (1), 2007

Galeano G., S. Suarez., H. Balslev. (1998). Vascular plant species count in a wet forest in the Choc area on the Pacific coast of Colombia. Biodiversity and Conservation 7: 1563-1572. Garca-Barriga H. (1975). Flora medicinal de Colombia. Tomo II, Instituto de Ciencias Naturales, Universidad Nacional Bogot, D.E. Colombia. 253-264. Gelach G., T. Ulmer. (2000). Una nueva especie de Passiflora, subgnero Passiflora (Passifloraceae) del oeste de Colombia. Caldasia 22 (2): 231-235. Gentry A.H. (1976). Additional Panamanian Passifloraceae. Annals of the Missouri Botanical Garden 63: 341-345. Gentry A.H. (1986). Species richness and floristic composition of Choco Region plant communities. Caldasia 15: 7175. Gentry A.H. (1992). New species of woody plants from Amazonian Peru. Novon 2(4): 333-338. Gentry A.H. (1993). Riqueza de especies y composicin florstica. In: P. LEYVA (Ed.). Colombia Pacfico, Vol. 1. Fondo Proteccin del Medio Ambiente Jos Celestino Mutis, Publicaciones Financiera Elctrica Nacional (FEN), Bogot. Gilbert L.E. (1982). The evolution of a butterfly and a vine, Heliconious butterflies. Scientific American 247: 110-121. Hansen A.K., G. Lawrence., B.B. Simpson., S.R. Downie., R. Stephen., A.C. Cervi., R. K. Jansen. (2006). Phylogenetic relationships and chromosome number evolution in Passiflora. Systematic Botany 31(1): 138-150. Harms, H. (1925). Passifloraceae. In: Engler A, ed. Die Natrlichen Pflanzenfamilien. Leipzig: Wilhelm Engelmann. 470-507. Hernndez J., A. Hurtado, R. Ortiz, T. Walschburger. (1991). Unidades Biogeogrficas de Colombia En: Hernndez J., R. Ortiz, T. Walshburger, A. Hurtado (Eds.) Estado de la Biodiversidad en Colombia Informe Final Santaf de Bogot, Instituto Colombiano para el Desarrollo de la Ciencia y la Tecnologa Francisco Jos de Caldas - Conciencias. Hernndez A. (2003). Revisin taxonmica de Passiflora, subgnero Decaloba (Passifloraceae) en Colombia. Tesis, Facultad de Ciencias, Departamento de Biologa, Universidad Nacional de Colombia sede Bogot. 138pp. Hernndez A., R. Bernal. (2000). Lista de especies de

Ocampo et al.

Colombian Passifloraceae

-33

Passifloraceae de Colombia. Biota Colombiana 1(3): 320-335. Hernndez A., N. Garca (2006). Las pasifloras (familia Passifloraceae). En: Libro rojo de plantas de Colombia. Las bromelias, las labiadas y las pasifloras. Garcia N., and Galeano G. (eds.). Volumen 3: 553-663. Hijmans R.J., M. Schreuder, M. De la Cruz, E. Rojas. (2001). Computer tools for spatial analysis of plant genetic resources data: DIVA-GIS. Plant Genetic Resources Newsletter 27: 15-19. Hno. Daniel (J. Gonzlez Patio) (1968). Curiosidades de una flor y de una familia botnica (La Flor de la Pasin). Boletn del Instituto de la Salle, Bogot. 208: 261-270. Holm-Nielsen L.B. (1974). Notes on central Andean Passifloraceae. Botaniska Notiser 127:338-351. Holm-Nielsen L.B., P. M. Jrgensen., J.E. Lawesson. (1988). Passifloraceae. In: Harling & L. Andersson (eds.), Flora del Ecuador 31: 124pp. IUCN (2003). Guidelines for application of IUCN Red List criteria at regional levels: Version 3.0. IUCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge, UK. IUCN (2004). Guidelines for using the IUCN Red List categories and criteria. Standards and petitions subcommittee of the IUCN SSC Red List programme committee. IUCN, Gland, Switzerland and Cambridge, U.K. Jrgensen P.M., J.E. Lawesson., L.B. Holm-Nielsen. (1984). A guide to collecting passion flowers. Annals of the Missouri Botanical Garden 71(4): 1172-1174. Jrgensen P.M., C. Ulloa., J.E. Madsen., R. Valencia. (1995). A floristic analysis of the high Andes of Ecuador. Churchill, S. P., H. Balslev, E. Forero & J. L. Luteyn (eds.). Biodiversity and Conservation of Neotropical Montane Forests, Proceedings of the Neotropical Montane Forest Biodiversity and Conservation Symposium, The New York Botanical Garden, Bronx, NY: 221-237. Jrgensen P.M., S. Len-Ynez. (1999). Catalogue of the vascular plants of Ecuador. Monograph Systematic Botanic. Missouri Botanical Garden 75: iviii, 11182. Jussieu A.L. de. (1805). Second Mmoire sur la famille des Passiflores, et particulirement sur quelques

espces nouvelles du genre Tacsonia. Annales du Musum dHistoire Naturelle 6: 388- 396. Killip E.P., W.F. Meggers, D.F. Hewett. (1924). New species of Passiflora from Tropical America. Journal of the Washington Academy of Sciences 14 (5): 108-112. Killip E.P. (1930). Ten new species of Passiflora, mainly from Colombia and Peru. Journal of the Washington Academy of Sciences 20 (15): 374-381. Killip E.P. (1938). The American Species of Passifloraceae Field Museum of Natural History Publication, Botanical Series 19 (1, 2): 1-613. Killip, E.P. (1960). Supplemental notes to the American species of Passiflora with descriptions of new species. Contributions from the U.S. National Herbarium 35: 2 (Tomo I). Lemarck J.B. Monenet de. (1789). Encyclopdie Mthodique, Botanique 3(1). Paris: Panckoucke. Linneaus C. (1753). Species Plantarum. Vol. 2. Stockholm: L. Salvius. MacDougal J.M. (1992). New species of Passiflora subg. Plectostemma (Passifloaceae). Novon 2(4): 358367. MacDougal J.M. (1994). Revision of Passiflora section Decaloba, Pseudodysosmia (Passifloraceae). Systematic Botany Monographs 14: 146pp. MacDougal J.M. (2006). Passiflora sandrae (Passifloraceae), a new species from Panama. Novon 16: 85-88. MacNeely J.A., K.R. Miller., N.A. Reid., R.A. Mittemer., T.B. Wainer. (1990). Conserving the worlds biological diversity World Conservation Union, World Resources Institute, World Wildlife Fund U.S. World Bank, Washigton, D.C. Martin F.W., H.Y. Nakasone. (1970). The edible species of Passiflora. Economic Botany 24 (3): 333-343. Martin P.S., C.R. Szuter. (1999). War zones and game sinks in Lewis and Clarks West. Conservation Biology 13: 36-45. Master M.T. (1872). Passifloraceae. In Flora Brasiliensis 13 (1). Ed. K. Mrtius and A. Eichler. 529-628. May R.M. (1992). How many species inhabit the Earth? Scientific American 267(4): 18-24. Maxted N., Mabuza-Dlamini P., Moss H., Padulosi S.,
Biota Colombiana 8 (1), 2007

34-

Colombian Passifloraceae Jarvis A., Guarino L., (2004) African Vigna : An Ecogeographic Study, IPGRI, Rome, Italy.

Ocampo et al.

do IUCN red list criteria conflate rarity and threat?. Biodiversity and Conservation 15: 1903-1912. Roemer, M.J. (1846). Familiarum naturalium regni vegetabilis synopses monographicae. Landes-IndustrieComptoir, Weimar. Romero-C.R. (1956). Plantas de valor comercial del gnero Passiflora: Granadilla, curuba, badea, y otras. Agricultura Tropical 12 (6): 403-407. Romero-C.R. (1991). Frutas silvestres de Colombia. Segunda edicin actualizada, Volumen con 207 ilustraciones. Instituto Colombiano de Cultura Hispnica. 423-429 p. Segura S., G. Coppens dEeckenbrugge., L. Lpez., M. Grum., L. Guarino. (2003). Mapping the potential distribution of five species of Passiflora in Andean countries. Genetic Resources and Crop Evolution 50: 555-566. Silverstone-Sopkin P.A., J.E. Ramos-Prez. (1995). Floristic exploration and phytogeography of the cerro del Torr, Choc, Colombia. Biodiversity of Conservation of Neotropical Montane Forests: 16986. In Biodiversity and Conservation of Neotropical Montane Forests, Proceedings of a Symposium, New York Botanical Garden, 21-26 June 1993 (S.P. Churchill, H. Balslev, E. Forero and J.L. Luteyn, eds.). New York Botanical Garden, New York. Simpson B. (1975). Pleistocene changes in the flora of high tropical Andes. Paleobiology 1:273-294. Svenning J.C. (1998). The effect of land-use on the local distribution of palm species in an Andean rain forest fragment in northwestern Ecuador. Biodiversity and Conservation 7: 15291537. Tillet S. (2003). Passifloraceae. In: Flora of the Venezuelan Guayana Volume 7: Myrtaceae-Plumbaginaceae by Julian A. Steyermark, Paul E. Berry, Kay Yatskievych, and Bruce K. Holtz (eds.): 625-667. Triana, M., J.E. Planchon. (1873). Passifloraeae. Annales des Sciences Naturelles 17: 121-186. Ulmer T. (1999). Passiflora formosa sp. nov., a hitherto misunderstood taxon in Passiflora subgenus Tacsonia (Passifloraceae) from Colombia. Edinburgh Journal of Botany 56 (2): 195-198. Ulmer T., J.M. MacDougal (2004). Passiflora: Passionflowers of the world. Timber Press, Inc. 430pp. UNEP-WCMC. (2004). World Conservation Monitoring Centre of the United Nations Environment

Muschner V.C., A.P. Lorenz., A.C. Cervi., S.L. Boniato., T.T. Souza-Chies., F.M. Salzano., L.B. Freitas. (2003). A first molecular phylogenetic analysis of Passiflora (Passifloraceae). American Journal of Botany 90 (8): 1229-1238. Myers J.E. (2000). The biodiversity challenge: Expanded hot spots analysis. The Environmentalist 10: 243256. Olson D.M. E. Dinerstein. (1998). The Global 200: A representation approach to conserving the earths most biologically valuable ecoregions. Conservation Biology 12: 502515. Prez E. (1956). Plantas tiles de Colombia. Tercera redaccin muy corregida y aumentada, con XLV laminas en negro, otras en color y 752 figuras en el texto. Librera Colombiana Camacho Roldn (Cia, Ltda.) Bogot. 611- 614. Plukenet, L. (1696). Almagestum botanicum sive Phytographi Pluckenetianae onomasticon methodo synthetic digestum, exhibens stirpium exoticarum, rariorum, novarumque nomina, qu descriptionis locum supplere possunt. Selbstverlag, London. Porter-Utley K.E. (2003). Revision of Passiflora subgenus Decaloba Supersection Cieca (Passifloraceae). Thesis Ph.D. Florida of University. 444p. Radhamani T.R., L. Sudarshana., R. Krishnan. (1995). Defence and carnivory: dual roles of bracts in Passiflora foetida L. Journal of Biosciences 20: 657-664. Rangel J.O. (1995). La diversidad florstica en el espacio andino de Colombia. En S.P. Churchill et al., eds. Churchill, S. P., H. Balslev, E. Forero & J. L. Luteyn (eds.). 1995. Biodiversity and Conservation of Neotropical Montane Forests, Proceedings of the Neotropical Montane Forest Biodiversity and Conservation Symposium, The New York Botanical Garden, Bronx, NY: 187-205. Rangel J.O. (2002). El estado actual del conocimiento de la flora de Colombia. Pg. 570 en: Rangel J.O., J. Aguirre-C & M.G. Andrade-C. (eds), Libro de resmenes octavo congreso latinoamericano y segundo Colombiano de botnica Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogot. Robbirt K.M., D.L. Roberts., J.A. Hawkins. (2006). Comparing IUCN and probabilistic assessments of threat:
Biota Colombiana 8 (1), 2007

Ocampo et al.

Colombian Passifloraceae

-35

Programme. Species Data (unpublished, September 2004). Web site at: [Link] Cambridge, England: UNEP-WCMC. Uribe L. (1954). Dos nuevas Passifloraceae colombianas. Mutisia 21: 1-5. Uribe L. (1955a). Pasiflorceas y Begoniceas de la Real Expedicin Botnica del Nuevo Reino de Granada. Ediciones Cultura Hispnica. Madrid 26:1-98. Uribe L. (1955b). Sertula Florae Colombiana 2. Caldasia 7(32):159-165. Uribe L. (1957). Una nueva e interesante Passiflora de Colombia. Caldasia 7(35):335-338. Uribe L. (1958). Sertula Florae Colombiae 4. Caldasia 8 (37):127-130. Uribe L. (1972). Catalogo ilustrado de las plantas de Cundinamarca: Passifloraceae, Begoniaceae,

Melastomataceae. Instituto de Ciencias Naturales, Facultad de Ciencias, Universidad Nacional. (5): 11-41. Uribe L. (1977). Sertula Florae Colombiae 14. Caldasia 12 (56): 13-18. Vanderplank J. (2000). Passion Flowers. 3nd ed. The MIT Press. Cambridge, Massachusetts. 224 pp. Vanderplank J. (2006). Passiflora miniata: Passifloraceae. Curtiss Botanical Magazine 23 (3): 223-230. World Press Review. (1993). Colombias vanishing forests, World Press Review, June 1993, Vol. 40 (6): 43pp. Yockteng R., S. Nadot. (2004). Phylogenetic relationships among Passiflora species based on the glutamine synthetase nuclear gene expressed in chloroplast (ncpGS). Molecular Phylogenetics and Evolution 31: 379-396.

Appendix 1. Synonymy = valid name

Cieca auriculata M. Roemer, 1846 = Passiflora auriculata Kunth, 1817 Cieca coriacea (Juss) M. Roemer, 1846 = Pasiflora coriacea Juss., 1805 Cieca discolor M. Roemer, 1846 = Passiflora misera Kunth., 1817 Cieca pallida (L.) M. Roemer, 1846 = Passiflora pallida L., 1753 Decaloba alnifolia M. Roemer, 1846 = Pasiflora alnifolia Kunth, 1817 Decaloba biflora (Lam.) M. Roemer., 1846 = Pasiflora biflora Lam., 1789 Decaloba bogotensis (Benth.) M. Roemer, 1846 = Passiflora bogotensis Benth., 1845 Decaloba cuneata M. Roemer., 1846 = Passiflora cuneata Willd., 1809 Decaloba filipes M. Roemer, 1846 = Pasiflora filipes Benth, 1843 Decaloba holosericea M. Roemer, 1846 = Passiflora holosericea L., 1753 Decaloba jacquini M. Roemer, 1846 = Passiflora pulchella Kunth, 1817 Dilkea johannesii var. parvifolia Hoehne, 1915 = Dilkea johannesii [Link]., 1885 Dilkea acuminata Mast., 1871 = Dilkea retusa Mast., 1871 Dilkea magnifica Steyerm., 1968 = Dilkea retusa Mast., 1871 Dilkea wallisii Mast., 1872 = Dilkea retusa Mast., 1871 Disemma hahnii E. Fourn., 1869 = Passiflora hahnii Mast., 1872 Disemma hahnii Fourn., 1869 = Passiflora hahnii (Fourn.) Mast., 1872
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Distephana cuneata M. Roemer, 1846 = Passiflora bicuspidata ([Link].) Mast., 1872 Distephana spinosa (Poeep. & Endl.) M. Roemer, 1835 = Passiflora spinosa (Poepp. & Ende.) Mast., 1871 Granadilla rubra Moench, 1802 = Passiflora rubra L., 1753 Grandilla vespertilio Moench, 1802 = Passiflora vespertilio L., 1753 Passiflora erubescens Triana & Planch., 1873 = Passiflora erytrophylla Mast., 1872 Passiflora velata Mast., 1872 = Passiflora serrulata Jacq., 1767 Passiflora williamsii Killip, 1922 = Passiflora platyloba var. williamsii (Killip) A.H. Gentry, 1976 Passiflora adenophylla Mast., 1872 = Passiflora subpeltata Ortega, 1798 Passiflora alba Link & Otto, 1798 = Passiflora subpeltata Ortega, 1798 Passiflora albicans L. Uribe, 1958 = Passiflora uribei L. K. Escobar, 1988 Passiflora angustifolia Swartz, 1788 = Passiflora suberosa L., 1753 Passiflora appendiculata G.F.W. Mey., 1818 = Pasiflora auriculata Kunth, 1817 Passiflora bauhinifolia Kunth, 1817 = Passiflora alnifolia Kunth, 1817 Passiflora bifurca Mast., 1873 = Passiflora cuneata Willd., 1809 Passiflora bilobata Vell., 1827 = Passiflora rubra L., 1735 Passiflora boyacana Killip, 1960 = Passiflora crispolanata L. Uribe, 1954 Passiflora capsularis var. geminifolia DC., 1828 = Passiflora sexflora Juss., 1805 Passiflora caucaense Holm-Niels., 1974 = Passiflora emarginata Humb. & Bonpl., 1813 Passiflora chilensis Miers, 1826 = Passiflora pinnatistipula Cav., 1799 Passiflora cisnana Harms, 1894 = Passiflora rubra L., 1753 Passiflora corumbaensis Barb., 1898 = Passiflora cincinnata Mast., 1868 Passiflora cualiflora Harms, 1906 = Passiflora citrifolia (Juss.) Mast., 1871 Passiflora difformis Kunth, 1817 = Passiflora coriaceae Juss., 1805 Passiflora digitata L., 1763 = Passiflora serratodigitata L., 1753 Passiflora elegans Triana & Planch., 1873 = Passiflora quindiensis Killip, 1938 Passiflora emiliae Sacco, 1966 = Passiflora ambigua Hemsl. ex Hook., 1902 Passiflora eminula Mast., 1883 = Passiflora costata Mast., 1872 Passiflora eriocaula Harms, 1922 = Passiflora rugosa (Mast.) Triana & [Link]. rugosa,1873 Passiflora erosa Rusby, 1907 = Passiflora morifolia Mast., 1872 Passiflora erosa Rusby, 1906 = Passiflora morifolia Mast., 1872 Passiflora fulgens Wallis ex Morren, 1866 = Passiflora coccinea Aubl., 1775 Passiflora gigantifolia Harms, 1894 = Passiflora macrophylla Spruce ex Mast., 1883
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Passiflora glauca Humb. & Bonpl., 1813 = Passiflora arborea Spreng., 1826 Passiflora goudotiana Triana & Planch., 1873 = Passiflora cumbalensis (H. Karst.) Harms var. goudotiana (Triana & Planch.) L. K. Escobar, 1987 Passiflora heydei Killip, 1922 = Passiflora morifolia Mast., 1872 Passiflora hydrophila Barb Rodr., 1891 = Passiflora costata Mast., 1872 Passiflora incana Seemann ex Mast., 1883 = Passiflora seemanni Griseb., 1858 Passiflora inundata Ducke, 1925 = Passiflora costata Mast., 1872 Passiflora laticualis Killip, 1924 = Passiflora misera Kunth., 1817 Passiflora longipes var. retusa Triana & Planch., 1873 = Passiflora longipes Juss., 1805 Passiflora lorifera Mast. & Andr, 1883 = Passiflora macrophylla Spruce ex Mast., 1883 Passiflora lunata J.E. Smith., 1790 = Passiflora biflora Lam., 1879 Passiflora macrocaropa Mast., 1869 = Passiflora quadrangularis l., 1759 Passiflora micrantha Killip, 1938 = Passiflora erythrophylla Mast., 1872 Passiflora miraflorensis Killip, 1924 = Passiflora sexflora Juss., 1805 Passiflora mollis var. integrifolia Planch. ex Mast., 1872 = Passiflora cuspidifolia Harms, 1893 Passiflora nympheoides Karst., 1859 = Passiflora nitida Kunth, 1817 Passiflora oblongifolia Pulle, 1906 = Passiflora laurifolia L., 1753 Passiflora ocanensis Planch. & Linden, 1873 = Passiflora lindeniana Planch. ex Triana & Planch., 1873 Passiflora ornata Kunth, 1817 = Passiflora maliformis L., 1753 Passiflora pala Planch. & Linden, 1873 = Passiflora bogotensis Benth., 1845 Passiflora paraguayensis Chad., 1899 = Passiflora capsularis L., 1753 Passiflora pennipes Sm., 1819 = Passiflora pinnatistipula Cav., 1799 Passiflora praeacuta Mast., 1887 = Passiflora oerstedii Mast., 1872 Passiflora pubera Planch. & Linden, 1873 = Passiflora sphaerocarpa Triana & Planch., 1873 Passiflora pulchella Kunth,1817 = Passiflora bicornis Mill., 1768 Passiflora quadriglandulosa var. involucrata (Mast.) Killip, 1938 = Passiflora involucrata (Mast.) A.H. Gentry, 1981 Passiflora reticulata Sauv., 1873 = Passiflora holosericea L.,1753 Passiflora salmonea Harms, 1894 = Passiflora parritae (Mast.) Bailey, 1916 Passiflora sanguinea J.E. Smithi, 1819 = Passiflora vitifolia Kunth, 1817 Passiflora schultzei Harms, 1929 = Passiflora arborea Spreng., 1826 Passiflora spherocarpa var. pilosula Mast., 1883 = Passiflora pubera Planch. & Linden, 1873 Passiflora stipulata Aubl., 1858 = Passiflora subpeltata Ortega, 1798 Passiflora suberosa var. pallida (L.) Mast. = Passiflora pallida L., 1753
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Passiflora tomentosa Lam. var. mollissima Triana & Planch., 1873 = Passiflora mollissima (Kunth) L.H. Bailey, 1916 Passiflora trisecta Planch. & Linden ex Triana & Planch., 1873 = Passiflora trianae Killip, 1938 Passiflora Van-Volxemii Triana & Planch., 1893 = Passiflora antioquienesis Karst., 1859 Passiflora var. cuellensis Goudot ex Triana & Planch., 1873 = Passiflora menispermifolia Kunth, 1817 Passiflora vesicaria L., 1753 = Passiflora foetida L., 1753 Passiflora vitifolia var. involucrata Mast., 1872 = Passiflora involucrata (Mast.) A.H. Gentry, 1981 Passiflora weberiana Andr, 1885 = Passiflora morifolia Mast., 1872 Passiiflora acerifolia Schlecht. & Cham., 1830 = Passiflora adenopoda Moc. & Sess ex DC., 1828 Rathea floribunda Karst., 1859 = Passiflora andina Killip, 1938 Tacsonia adulterina Juss., 1805 = Passiflora adulterina L. f., 1781 Tacsonia bicuspidata H. Karst., 1859 = Passiflora bicuspidata (H. Karst.) Mast., 1872 Tacsonia cumbalensis H. Karst., 1859 = Passiflora cumbalensis var. cumbalensis (H. Karst.) Harms, 1894 Tacsonia cuneata Benth, 1845 = Passiflora bicuspidata (H. Karst.) Mast.,1872 Tacsonia flexipes (Triana & Planch) Mast., 1883 = Passiflora flexipes Triana & Planch., 1873 Tacsonia glandulosa Juss., 1805 = Passiflora glandulosa Cav.,1790 Tacsonia infundibularis Mast., 1883 = Passiflora bracteosa Planch. & Linden, 1873 Tacsonia lanata Juss., 1811= Passiflora lanata (Juss.) Poir., 1811 Tacsonia mixta (L.f.) Juss., 1805 = Passiflora mixta L.f., 1781 Tacsonia mollissima Kunth var. glabrescens Mast.,1872 = Passiflora mollissima (Kunth) L.H. Bailey, 1916 Tacsonia mollissima Kunth, 1817 = Passiflora mollissima (Kunth) L.H. Bailey, 1916 Tacsonia parritae Mast., 1882 = Passiflora parritae (Mast.) L.H. Bailey, 1916 Tacsonia pinnatistipula var. pennipes (Sm.) DC., 1828 = Passiflora pinnatistipula Cav., 1799 Tacsonia pinnatistipula (Cav.) Juss., 1805 = Passiflora pinnatistipula Cav.,1799 Tacsonia quadriglandulosa (Rodschied) DC., 1828 = Passiflora quadriglandulosa Rodschied, 1796 Tacsonia rosea (H. Karst.) Sodiro, 1903 = Passiflora x rosea (H. Karst.) Killip, 1938 Tacsonia rugosa Mast., 1872 = Passiflora rugosa (Mast.) Triana & Planch, 1873 var. rugosa Tacsonia spinescens Klotsch in Schomb., 1848 = Passiflora securiclata Mast., 1893 Tacsonia spinosa Poepp. & Endl., 1835 = Passiflora spinosa (Poepp. & Ende.) Mast., 1871 Tacsonia trinervia Juss., 1805 = Passiflora trinervia (Juss.) Poir., 1811 Tetrastylis lobata Killip, 1926 = Passiflora lobata (Killip) Hutch. ex J.M. MacDougal, 1986

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Appendix 2. Total number of Passifloraceae present in Colombia.

Number of observations, Maximum distance (MaxD) and Circular area (CA) for each species. Endemic species are highlighted by an asterisk (*). RC: species rare for Colombia; Roc: species rare in other countries; Rne: rare narrow endemic, Ne: narrow endemic; Re: rare endemic; Ce: common endemic.
Species Ancistrothyrsus antioquiensis L.K. Escobar (ined.)* Ancistrothyrsus tessmannii Harms Dilkea johannesii Barb. Rodr. Dilkea parviflora Killip Dilkea retusa Mast. Passiflora adenopoda Moc. & Sess ex DC. Passiflora adulterina L.f. * Passiflora alnifolia Kunth Passiflora alata Curtis Passiflora ambigua Hemsl. ex Hook. Passiflora andina Killip Passiflora andreana Mast. Passiflora antioquiensis H. Karst. * Passiflora apoda Harms Passiflora arbelaezii L. Uribe Passiflora arborea Spreng. Passiflora auriculata Kunth Passiflora azeroana L. Uribe * Passiflora bicornis Mill., 1768 Passiflora bicuspidata (H. Karst.) Mast. * Passiflora biflora Lam. Passiflora bogotensis Benth. * Passiflora bracteosa Planch. & Linden Nb. observ. 2 1 1 22 5 51 43 121 1 48 1 3 55 43 48 67 128 10 11 16 40 56 7 MaxD (km) 41 0 0 1,185 952 383 234 1,244 0 929 0 45 667 678 746 1,204 1,635 574 675 438 1,326 1,057 122 CA (km2) 11,762 7,814 7,814 40,688 106,159 82,65 39,072 170,761 7,814 137,261 7,814 12,214 99,064 83,615 113,491 144,115 334,952 34,734 52,098 61,674 122,047 89,25 23,18 RC /Roc Ce Colombia and Venezuela Ce Ce RC /Roc RC Ce Colombia and Ecuador RC Ce RC Rare species RC RC / Roc RC / Roc Endemics and distribution Rne (Antioquia)

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Species Passiflora bucaramangensis Killip * Passiflora callistema L.K. Escobar * Passiflora candollei Tr. & Planch. Passiflora capsularis L. Passiflora chelidonea Mast. Passiflora chocoensis G. Gerlach & T. Ulmer * Passiflora cincinnata Mast. Passiflora citrifolia (Juss.) Mast. Passiflora coccinea Aubl. Passiflora colombiana L.K. Escobar * Passiflora coriacea Juss. Passiflora costaricensis Killip Passiflora cremastantha Harms * Passiflora crispolanata [Link] * Passiflora cuatrecasasii Killip * Passiflora cumbalensis (Karst.) Harms Passiflora cuneata Willd. Passiflora cuspidifolia Harms, 1893 Passiflora danielii Killip * Passiflora dawei Killip * Passiflora emarginata Humb. & Bonpl.* Passiflora engleriana Harms * Passiflora erytrophylla Mast. * Passiflora escobariana J.M. MacDougal Passiflora filipes Benth. Passiflora fimbriatistipula Harms * Passiflora flexipes Triana & Planch. *

Nb. observ. 8 1 4 64 18 1 1 3 21 2 59 1 1 11 9 156 9 33 5 4 46 2 6 2 3 18 24

MaxD (km) 70 0 854 1,437 1,024 0 0 68 1,285 42 741 0 0 246 181 1,196 877 812 180 208 654 110 225 3 48 198 322

CA (km2) 15,032 7,814 26,294 159,962 94,209 7,814 7,814 14,049 107,128 11,91 136,372 7,814 7,814 29,72 21,312 199,941 50,607 86,64 20,59 23,702 78,393 8,902 27,643 8,136 13,227 33,664 36,121

Rare species RC RC RC

Endemics and distribution Ne (Santander) Rne (Bolivar)

RC RC RC

Rne (Choco)

RC

Rne

RC RC Rne (Cauca) Ce Ne

RC RC

Rne (Antioquia) Rne Ce

RC

Rne (Antioquia) Ne

RC RC / Roc Mxico to Ecuador Ne Ce

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Species Passiflora foetida L. Passiflora formosa T. Ulmer * Passiflora glandulosa Cav. Passiflora gleasonii Killip Passiflora gracillima Killip Passiflora grandis Killip * Passiflora gritensis H. Karst. Passiflora guatemalensis S. Watson Passiflora guazumaefolia Juss. Passiflora hahnii (Fourn.) Mast. Passiflora haughtii Killip * Passiflora hirtiflora Jorgensen & Holm-Nielsen Passiflora holosericea L. Passiflora holtii Killip Passiflora hyacinthiflora Planch. & Linden * Passiflora involucrata (Mast) A.H. Gentry Passiflora jardinensis L.K. Escobar * Passiflora kalbreyeri Mast. * Passiflora killipiana Cuatrecasas Passiflora lanata (Juss.) Poir. * Passiflora laurifolia L. Passiflora lehmanni Mast. * Passiflora leptomischa Harms * Passiflora ligularis Juss. Passiflora lindeniana Planch. ex Triana & Planch. Passiflora linearistipula L.K. Escobar *

Nb. observ. 143 1 1 2 29 2 8 11 8 1 1 1 7 1 3 8 8 19 1 32 11 17 21 101 2 4

MaxD (km) 1,83 0 0 3 684 14 346 971 349 0 0 0 238 0 305 1,197 35 426 0 284 1,35 805 449 914 395 8

CA (km2) 420,44 7,814 7,814 8,075 74,546 9,161 26,115 59,505 41,192 7,814 7,814 7,814 25,632 7,814 17,746 48,827 11,335 41,237 7,814 45,476 84,672 91,156 46,331 170,123 15,628 8,695

Rare species

Endemics and distribution

RC RC RC

Rne (Boyac)

RC

Rne

RC RC RC Rne (Santander)

RC / Roc RC Re

RC

Ne (Antioquia) Ce

RC / Roc

Colombia to Peru Ce

Ce Ce

RC / Roc RC

Colombia and Venezuela Rne (Caldas)

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Species Passiflora lobata (Killip) Hutch. ex J.M. MacDougal Passiflora longipes Juss. * Passiflora lyra Planch. & Lind. ex Killip Passiflora macrophylla Spruce ex Mast. Passiflora magdalenae Triana & Planch. * Passiflora magnifica L.K. Escobar* Passiflora maliformis L. Passiflora manicata (Juss.) Pers. Passiflora mariquitensis Mutis ex Uribe * Passiflora megacoriacea Porter-Utley (ined.) Passiflora menispermacea Triana & Planch. * Passiflora menispermifolia Kunth Passiflora micropetala Mast. Passiflora misera Kunth Passiflora mixta L. f. Passiflora mollis Kunth Passiflora monadelpha Jorgensen & Holm-Nielsen Passiflora morifolia Mast. Passiflora multiformis Jacq. Passiflora munchiquensis Hernandez (ined.)* Passiflora mutisii Killip * Passiflora nitida Kunth Passiflora occidentalis Hernandez (ined.)* Passiflora oerstedii Mast. Passiflora pacifica L.K. Escobar * Passiflora palenquensis Holm-Niels. & Lawesson Passiflora pamplonensis Planch.& Linden ex Tr. & Planch. *

Nb. observ. 3 21 4 20 21 6 122 62 3 1 2 43 11 54 162 17 7 1 4 4 1 72 10 41 9 20 1

MaxD (km) 194 334 69 716 129 33 1,208 889 10 0 18 1,41 1,318 1,148 966 554 67 0 147 200 0 1,452 474 728 510 1,181 0

CA (km2) 23,115 45,557 14,716 90,432 31,127 12,215 212,27 114,036 8,436 7,814 9,61 167,659 68,015 145,398 191,787 208,941 33,665 7,814 17,652 22,441 7,814 279,511 42,35 148,975 39,585 100,769 7,814

Rare species RC

Endemics and distribution

Ce RC / Roc

Ne RC Ne (Antioquia)

RC RC RC

Rne (Tolima)

Rne (Tolima)

RC / Roc RC RC RC RC

Colombia and Ecuador

Rne Rne (Tolima)

Ce

Ce

RC

Rne (N. de Santander)

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Species Passiflora pallida L. Passiflora panamensis Killip Passiflora parritae (Mast.) L.H. Bailey * Passiflora pennellii Killip * Passiflora phaeocaula Killip Passiflora picturata Ker Passiflora pilosissima Killip * Passiflora pinnatistipula Cav. Passiflora pittieri Mast. Passiflora platyloba Killip Passiflora popayanensis Killip * Passiflora popenovii Killip Passiflora punctata L. Passiflora purdiei Killip * Passiflora putumayensis Killip Passiflora pyrrhantha Harms Passiflora quadrangularis L. Passiflora quadriglandulosa Rodschied Passiflora quindiensis Killip * Passiflora resticulata Mast. & Andr Passiflora rigidifolia Killip * Passiflora riparia Mart. ex Mast. Passiflora rubra L. Passiflora rugosa (Mast.) Triana & Planch Passiflora schlimiana Triana & Planch. * Passiflora securiclata Mast. Passiflora seemannii Griseb.

Nb. observ. 6 15 14 6 5 1 2 21 1 4 6 17 8 1 1 1 112 4 8 4 1 3 90 12 7 4 40

MaxD (km) 898 295 100 313 498 0 270 750 0 201 64 636 592 0 0 0 1,676 414 225 414 0 716 1,351 421 181 849 1,341

CA (km2) 50,078 41,614 20,357 24,413 28,305 7,814 15,628 57,114 12,661 16,471 15,078 31,075 40,022 7,814 7,814 7,814 314,317 21,256 24,711 18,938 7,814 23,442 117,934 35,549 27,852 30,708 129,777

Rare species

Endemics and distribution

RC

Ne Ce

RC / Roc RC Re

RC RC RC Ne (Cauca)

RC RC / Roc RC / Roc

Rne

RC Ne (Tolima)

RC RC

Rne (Antioquia)

Roc RC / Roc

Ne Colombia and Venezuela

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Species Passiflora semiciliosa Planch & Linden * Passiflora serratodigitata L. Passiflora serrulata Jacq. Passiflora sexflora Juss. Passiflora sierrae L.K. Escobar * Passiflora skiantha Huber Passiflora smithii Killip Passiflora sodiroi Harms Passiflora sphaerocarpa Triana & Planch. * Passiflora spicata Mast. Passiflora spinosa (Poepp. & Endl.) Mast. Passiflora suberosa L. Passiflora subpeltata Ortega Passiflora tarminiana Coppens & Barney Passiflora tenerifensis L.K. Escobar * Passiflora tica Gmez-Laur. & L.D. Gmez Passiflora tiliifolia L. Passiflora tolimana Harms * Passiflora trianae Killip * Passiflora tribolophylla Harms * Passiflora tricuspis Mast. Passiflora trinervia (Juss.) Poir.* Passiflora tripartita (Juss.) Poir. Passiflora trisulca Mast. * Passiflora truxillensis Planch. & Linden ex Triana & Planch. Passiflora tryphostemmatoides Harms Passiflora tuberosa Jacq.

Nb. observ. 4 18 10 14 2 1 28 1 35 1 20 66 35 28 4 5 48 12 2 1 1 27 56 8 1 25 1

MaxD (km) 578 1,566 331 353 46 0 827 0 878 0 1,521 1,497 1,344 832 71 319 1,01 426 39 0 0 220 1,21 441 0 557 0

CA (km2) 26,175 67,105 29,354 43,143 12,194 7,814 72,555 7,814 96,244 7,814 118,197 158,86 89,527 103,373 15,195 23,119 97,205 33,711 11,594 7,814 7,814 36,932 145,398 25,258 15,628 77,831 7,814

Rare species RC

Endemics and distribution Re

RC RC / Roc

Rne (Magdalena) Colombia and Peru

RC / Roc

Colombia and Ecuador Ce

RC / Roc

Colombia and Brazil

RC RC

Rne (Valle del Cauca)

Ce RC RC RC Ne Rne Rne

Ce RC / Roc Colombia and Venezuela

RC

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Species Passiflora uribei L.K. Escobar * Passiflora ursina Killip & Cuatrec. Passiflora variolata Poepp. & Endl. Passiflora venosa Rusby Passiflora vespertilio L. Passiflora vestita Killip Passiflora vitifolia Kunth Passiflora x rosea (H. Karst.) Killip

Nb. observ. 3 2 6 1 3 1 359 7

MaxD (km) 54 7 412 0 292 0 1,729 161

CA (km2) 12,96 8,503 27,059 7,814 20,887 7,814 456,229 20,988

Rare species RC RC / Roc

Colombia and Ecuador

RC / Roc RC RC / Roc Colombia and Ecuador

P. emarginata

Recibido: 18/01/2007 Aceptado: 13/03/2007

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