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Una especie nueva de Castilleja (Orobanchaceae) de los paramos de la cordillera oriental de colombia, con comentarios acerca de su asociacion con Plantago rigida (Plantaginaceae).

A NEW SPECIES OF CASTILLEJA (OROBANCHACEAE) FROM THE PARAMOS OF THE COLOMBIAN EASTERN CORDILLERA, WITH COMMENTS ON ITS ASSOCIATION WITH PLANTAGO RIGIDA (PLANTAGINACEAE)

INTRODUCTION

Castilleja Mutis ex L. f. comprises approximately 200 species (Holmgren 1978, Chuang & Heckard 1991, Heide-J0rgensen 2008) most of which are native to western North America. The South American species (estimated between 15 and 20; Egger, pers. comm.) are concentrated in the Andes. During the ongoing study of parasitic and hemiparasitic plants in Colombia by the authors, a number of specimens of Castilleja collected in the paramos of the Eastern Cordillera drew our attention because of the small size of the individuals and the frequent association with Plantago rigida Kunth. A specimen recently collected by FG in the paramo of Siscunsi (Sogamoso, Boyaca) matches all the morphological and ecological traits of the small-sized individuals previously collected in similar paramos of Santander, northern Boyaca, and Cundinamarca, in the Colombian Eastern Cordillera. Until now, these specimens have remained either undeterminated or were tentatively identified as infraspecific taxa of C. fissifolia L. f. or assigned to C. nubigena Kunth, a species known from the high Andes of Ecuador and Peru (cf. Holmgren 1984, Luteyn 1999, Molau, 1999). A detailed examination of these collections reveals, however, that they do not match any of the previously described species of Castilleja from Costa Rica/Panama (cf. Holmgren 1978) Venezuela (cf. Pennell 1953, Egger 2009), Colombia (cf. Fernandez-Alonso 1987), Ecuador (cf. Holmgren 1984) or Peru (cf. Weddell 1857, Herrera 1926, Edwin, 1971). Thus, here we describe the new species and compare it with other high Andean species of Castilleja characterized by the small size of the individuals. Besides the disjunct distribution between the new species and the other morphologically similar high Andean species (C. nubigena, C. pumila (Benth.) Wedd. ex Herrera, and C. virgata (Wedd.) Edwin), the newly described species possesses clear-cut differences in the size and overall morphology of leaves, bracts, and flowers (Table 1).

Additionally, we include preliminary observations on the association of the species with Plantago rigida, as root hemiparasitism and hence physical plant interactions may also have genetic implications, for instance the occurrence of Horizontal Gene Transfer (HGT).

HGT has been reported between members of the Orobanchaceae and Plantaginaceae, preferentially involving mitochondrial and ribosomal over plastid genes (Mower et al. 2004, Davis & Wurdack 2004). For instance, the mitochondrial atp1 has been transferred from Bartsia (Orobanchaceae) to Plantago (Mower et al. 2004). Plastid HGT seemed to have occurred less frequently, like between the parasitic Orobanche and Pelipanche (Park et al. 2007). These are both non-photosynthetic parasites with a shared plastid region including rps2, trnL-F, and rbcL that has probably been transferred via their common host (Park et al. 2007). At this point the genetic consequences of the Castilleja paramensis-Plantago rigida association are still unknown; however, their tight physical link and their exclusive distributions in high altitudes in the Andes, make them an excellent candidate to undertake studies testing explicitly for the occurrence of HGT.

Castilleja paramensis F. Gonzalez & PabonMora, sp. nov. TYPE: COLOMBIA. Boyaca: Sogamoso, Paramo de Siscunsi, alrededores de la Laguna de Siscunsi, 3875 m, 5[degrees]22'N, 72[degrees]50'W, 04 Feb 2013 (fl), F. Gonzalez, G. Plunkett, A. Nicolas, J. Aguirre & A. Herrera 4478 (Holotype: COL; isotypes: HUA, NY, UPTC, to be distributed). Figs. 1-3

Small, rhizomatous, decumbent, perennial herbs; ascending stems to 5(-8) cm tall, branching from the base. Rhizomes very thin, whitish, with a few scattered achlorophyllous triangular scale-leaves to 2 mm long and 1 mm wide. Internodes of the aerial shoots very short, to 3 mm long. Leaves congested, sub-opposite, cucullate, sessile, broadly elliptic, to 7 mm long and 5 mm wide, entire to slightly bi- or trilobate, light green when young, later with the distal half purplish, lateral lobe(s) triangular, ascending, glabrous above, scarcely villous below, margins villous, especially along the distal half. Bracts broadly elliptic, bi- or trilobate and bright red in their distal half, glabrous above, villous below, with those subtending the mature flowers becoming purplish. Flowers subsessile, pedicel < 1 mm long, axillary to the 3-4 most distal bracts. Calices 8-11 mm long, green when immature, with bright yellow distal half at maturity, not inflated, villous, tube to 6 mm long, median clefts strongly unequal, the adaxial 1-1.5 mm deep, the abaxial 4.55.5 mm deep, lateral clefts absent. Corollas 8-11.5 mm long, completely included or very slightly exserted from the calices, hyaline, straight to very slightly incurved, tubes 5-6 mm long, the galeas 4-5.5 mm long, villous along the adaxial mid-line, the lower three lips consisting of bright green, extremely reduced, triangular, to 0.5 mm long and 0.2 mm wide, ascending, glabrous. Stamens included, slightly didynamous; filaments 8.5-10.5 mm long, adnate to the corolla tube for its proximal 1/3, anthers lanceolate, ca. 0.5-0.8 mm long, thecae slightly asymmetric. Styles as long as the corollas and included during anthesis.

Stigmas slightly bilobed, poorly differentiated from the styles, papillose. Capsules oblong, to 10 x 5 mm, villous. Seeds obconical, 1.3-1.5 x 0.8-1 mm, seed coat papery, translucent, unruptured, with deep and narrowly hexagonal reticulation cells.

Etymology: The specific name refers to the ecosystem where the species has been found.

Distribution and phenology: Castilleja paramensis has so far been collected in paramos of the departments of Santander, Boyaca and Cundinamarca (Colombia). It sets flowers in September, November to March and in May, and fruits in February, June and September.

Vernacular name: "Flores de lagarto" (Guican, Boyaca; FG 1391).

Additional specimens examined (when available, we have included the field notes referring to the association of the new species with Plantago rigida). COLOMBIA. Boyaca: Morrena seca 200 m NW de la Laguna Pintada, con Espeletia colombiana predominante, asociada con Aciachne pulvinata, Agrostis breviculmis, Agrostis trichoides y Acaena cylindrostachya, 3995 m, 25 Sep 1972 (fl, fr), A. M. Cleef & P. A. Florschutz 5508 (COL); Paramo de La Rusia, NW-N de Duitama, Pena Blanca, 2 km al NE de Buenos Aires, pantanos con almohadas de Plantago rigida predominantes, 3720 m, 16 Dic 1972 (fl), A. M. Cleef 7317 (COL); Sierra Nevada del Cocuy, paramo Concavo, superparamo en el limite con el paramo p.d., 3.5 km al NNW del morro Pulpito del Diablo, cabeceras de vallecitos entre morrenas, protegidas a viento con un frailejonal denso y humedo de Espeletia lopezii, 4335 m, 26 Feb 1973 (fl, fr), A. M. Cleef 8530 (COL); Socota, paramo de Pisba, sitio La Playita, entre colchones de Plantago rigida (FG 824), c 3600 m, 15 Nov 1986 (fl), F. Gonzalez et al. 823 (UPTC); Socha, sitio Los Pinos, 3600 m, asociada a Plantago rigida (FG 1118A), 25 Jun 1987 (fl), F. Gonzalez et al. 1118 (UPTC); Guican, ascenso al Ritacuba Blanco, por las cabanas del Himat, 3800-4800 m, 1 Mar 1988 (fl), F. Gonzalez et al. 1391 (UPTC); Guican, Sierra Nevada del Cocuy, carreteable entre la cabana del Inderena y las Lagunillas, frente a la Laguna La Atravesada, 4000 m, entremezclada con Plantago rigida (FG 1417), 2 Mar 1988 (fl), F. Gonzalez et al. 1416 (UPTC); Belen, paramo del Guina, 3500 m, 22 May 1993 (fl), F. Gonzalez & R. Small 2720 (COL, UPTC); Sierra Nevada del Cocuy, sector SE, valle del rio Lagunillas, entre 3650 y 4000 m, 24 May 1993 (fl), F. Gonzalez & R. Small 2734 (COL, UPTC); Guican, ascenso hacia el Boqueron de Cardenillo, 6[degrees]18'50.6"N, 72[degrees]12'15.6"W, ca. 4300 m, 17 Ago 2013 (fl), S. Gonzalez et al. 01 (COL); Guican, Sierra Nevada del Cocuy, Laguna Grande de los Verdes, en colchones de Distichia sp., 3900-4100 m, 20 Sep 1978 (fl, fr), O. Rangel & H. Sturm 1534A (COL); below the Playas of Ritacuba in the Cocuy mountains above Guican, locally frequent in poorly drained damp paramo 3700-4500 m, particularly in damp flushes and poorly drained grassy hollows, 4200 m, 24 Jun 1984 (fl, fr), J. R. I. Wood 4458 (COL). Cundinamarca: Paramo de Sumapaz, Chisaca, pantano con cojines de Plantago rigida, 3680 m, 11 Dic 1971 (fl), A. M. Cleef 143 (COL); Macizo de Bogota, paramo de Chisaca, around the lakes at about 3620 m, 16 Sep 1961 (fl), J. Cuatrecasas & R. Jaramillo M. 25972 (COL). Santander: Cordillera Oriental, paramo del Almorzadero, above 3600 m, slope above cienaga, very wet, with hummocks of mosses, Plantago rigida, etc., 31 Dec 1959-1 Jan 1960 (fl), H. G. Barclay & P. Juajibioy 10369 (COL).

Castilleja paramensis is morphologically similar to C. pumila, C. virgata and especially to C. nubigena (Table 1). It clearly differs by its smaller size (up to 8 cm), broadly elliptic and entire to slighly trilobed leaves and bracts, reduced pedicel (<1mm) and flower (0.8- 1.15 cm) lenght, bright yellow and not inflated calyx, and corolla included or very slightly (for < 2 mm) exserted from the calyx (Fig. 1). Additional differences between C. paramensis and C. nubigena include the absence of lateral clefts of the calyx (Fig. 3B, D), the straight to very slightly incurved corolla, the included stamens and the bilobed, poorly differentiated stigma in the former, versus the presence of lateral clefts of the calyx, the strongly incurved corolla, the exserted stamens and the capitate stigma in the latter. Finally, the presence of included corolla, stamens, styles and stigmas (Figs. 1, 3) in C. paramensis strongly suggests the ocurrence of cleistogamy in this species.

On the systematics and life history of Castilleja: The systematics of Castilleja has been considered difficult, mainly due to polyploidy, lack of crossing barriers among species, and effective natural hybridization, all of which can drastically change the morphological traits used to distinguish species (Heckard 1964, Heckard & Chuang 1977, Chuang & Heckard 1991, Mathews & Lavin 1998, Hersch-Green & Cronn 2009, Clay et al. 2012, Hersch-Green 2012). At a suprageneric level, polyploidy has also led to differential sampling of paralogs (e.g. PHYA) in phylogenetic analyses, resulting in a polyphyletic Castilleja (Bennett & Mathews 2006).

The South American species of Castilleja fall into two different subgenera, Castilleja and Colacus. All the Colombian species belong to the subgenus Castilleja (sensu Chuang & Heckard 1991), as the flowers are narrow and tubular, the associated bracts, the calices and/or corollas are red, and the galea is at least three times longer than the lower three lips. All these traits have been associated with hummingbird, pollination in the genus (Pennell 1948, Chuang & Heckard 1991, Heide-J0rgensen 2008). Hummingbird pollination, in turn, appears to be correlated with the evolution of perenniality in the genus (Tank & Olmstead 2008, 2009). This perennial habit has evolved once from an annual condition in Castilleja, and it seems to be tightly associated with polyploidy at least in the North American species (Tank & Olmstead 2008). The new species can be assigned unequivocally to subgenus Castilleja, and it is morphologically and ecologically similar to the high-Andean species C. nubigena, C. pumila and C. virgata (Table 1). The latter three species form a clade of perennial but primarily diploid species (Tank & Olmstead 2009). We anticipate that the perennial C. paramensis belongs to this clade; however, chromosome counts and molecular data are needed to confirm this placement. In addition, studies on pollination biology of C. paramensis are critical, as its colorful flowers are near the ground and appear to be too small and too low for hummingbird pollination. The included style and stigma of C. paramensis (Fig. 3) strongly suggest that the species might be cleistogamous, despite the fact that it can co-occur with C. integrifolia, a species pollinated by the hummingbird Oxypogon guerinii (Salamanca-Reyes 2011).

Castilleja species as facultative root hemiparasites. Species of Castilleja are facultative root hemiparasites, which means they can complete their life cycle either as parasites or singly without a host (Heckard 1962, 1964, Matthies 1997, Heide-Jorgensen 2008, Fay et al. 2010). Castilleja host association is broad and can infect species across a diverse range of flowering plant families (Heckard 1962, Musselman & Press 1995), though some species might have more exclusive associations. This is the case for C. paramensis, frequently found associated belowground with Plantago rigida (Fig. 2). In fact, Cleef (1981:127) included C. paramensis (referred there as "C. fissifolia ssp. pygmaea, ssp. nom. herb.; in ed.?") and P. rigida as two primary components of the Wernerietea syntaxonomy that conforms the "flush vegetation and cushion bogs".

Seed germination in Castilleja can occur in the absence of a host (Heckard 1962); however, species from high elevations may require moist and cold conditions that can slow down germination rates (Kuijt 1969). Heckard (1962) examined hemiparasitic Castilleja species growing with and without hosts and noticed that generally plants growing as parasites are more vigorous, branch more profusely, and exhibit early flowering. Haustoria in Castilleja are small (less than 1 mm in diameter) and are not as invasive or penetrating as haustoria of other parasitic flowering plants. They initiate as elongated epidermal cells similar to root hairs (Kuijt 1969). The first responses during haustorial initiation are cortical enlargement and increasing cell division, resulting in lateral or apical protuberances in the parasite root similar to other Orobanchaceae, such as Bartsia, Lathraea, Melampyrum, Pedicularis and Rhinanthus (Heckard 1962, Dobbins & Kuijt 1973a, Weber & Weberling 1975, Riopel & Timko 1995). A mature haustorium of Castilleja is composed of a plate xylem zone surrounded by parenchyma, a centrally located core of collenchyma (with vessel members scattered), and the haustorial surface (Dobbins & Kuijt 1973a). Penetration is likely due to cell dissolution in the host, accompanied by crushing of cells (Dobbins & Kuijt 1973b). The haustoria of C. paramensis are spherical to cup-shaped and usually terminal in secondary roots.

ACKNOWLEDGEMENTS

Thanks to J. Aguirre, A. Nicolas, and G. Plunkett (The New York Botanical Garden), and A. Herrera and S. Gonzalez (National University of Colombia), for accurate observations and valuable help during fieldwork while collecting the type specimen and some of the paratypes. Thanks also to J. M. Egger (Herbarium, Burke Museum of Natural History and Culture, University of Washington, Seattle), O. Rangel (Institute of Natural Sciences, National University of Colombia), and three anonymous reviewers, for valuable comments on the manuscript.

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Recibido: 19/02/2013

Aceptado: 05/09/2013

FAVIO GONZALEZ

Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apartado 7495, Bogota, D.C., Colombia. fagonzalezg@unal.edu.co

NATALIA PABON-MORA

Instituto de Biologia, Universidad de Antioquia, Apartado 1226, Medellin, Colombia.

Table 1. Comparative traits of Castilleja paramensis and the three
high/Andean small perennial herbaceous species.

                        C. paramensis sp.      C. nubigena Kunth
                               nov.                   (1)

Habit                    decumbent herbs,       erect subshrubs,
                       with ascending stems    stems diffuse, 10-
                        to 5(-8) cm tall;     18.4 cm tall; leaves
                       leaves sub-opposite       alternate, not
                        and very congested         congested

Shape and size of      broadly elliptic, to        linear to
leaf blade                 0.7 x 0.5 cm       lanceolate, trifid,
                                              1.12.5 x 0.3-0.6 cm

Number and shape of      0-1 pair, broad       1-2 pairs, narrow
lateral lobes of the
leaf

Shape and color of      broadly elliptic,        lanceolate to
the bracts             entire or with 1 or    broadly lanceolate,
                         2 broad lateral       deeply 3-5-lobate,
                        lobes, bright red          red tipped
                         when young, then
                             purplish

Length of the                 <1 mm                  2-4 mm
pedicel

Total length of the        0.8-1.15 cm             1.55-2 cm
flower

Calyx versus corolla   corolla included or      corolla exserted
                        very slightly (for     from the calyx for
                       <2 mm) exserted from          >4 mm
                            the calyx

Color of the calyx       bright yellow at     red to occasionally
                           least in the             purplish
                          marginal zone

Shape and length of     not inflated, 8-11     inflated, 11-16 mm
the calyx                       mm                    long

Adaxial and abaxial     very unequal, the      very unequal, the
clefts of the calyx     adaxial 1-1.5 mm,     adaxial 1-3 mm deep,
                       the abaxial 4.5-5.5    the abaxial (6-)10-
                                mm                 14 mm deep

Lateral clefts of             absent            present, to 1 mm
the calyx                                             deep

Color of the corolla     uniformly bright           greenish
                              green

Length and shape of    8-11.5 mm, straight       10-15(-20) mm,
corolla                  to very slightly      strongly incurved
                             incurved          especially between
                                                the tube and the
                                                     lobes

Length of the            5-6 mm, usually            6-7 mm,
corolla tube             longer than the        approximately as
                              galea            long as the galea

Length of the galea          4-5.5 mm                4-9 mm

Stamens                 included style as           exserted
                           long as the

Style and stigma in     corolla, included.     exserted from the
anthesis                 Stigmas slightly     corolla and from the
                         bilobed, poorly         calyx; stigma
                       differentiated from          capitate
                           the styles,
                            papillose

Length of the seeds         1.3-1.5 mm             1.6-2.1 mm

Distribution and        Santander, Boyaca,       Azuay, Carchi,
altitudinal range       and Cundinamarca;         Chimborazo,
                        eastern slopes of       Cotopaxi, Napo,
                          the Colombian          Pichincha and
                       Eastern Cordillera.    Tungurahua (Ecuador;
                            35004335 m         endemic, according
                                                to Molau, 1999).
                                               Peru (according to
                                                 Luteyn, 1999).
                                               (3250-) 36004650 m

                        C. pumila (Benth.)     C. virgata (Wedd.)
                            Wedd. (2)              Edwin (3)

Habit                    decumbent herbs,        erect herbs or
                       stems to 9 cm tall;     subshrub, 20-40(-
                        leaves alternate,     60) cm tall; leaves
                          not congested          alternate, not
                                                   congested

Shape and size of         lanceolate to            linear to
leaf blade             broadly lanceolate,    lanceolate, 1.5-3(-
                        0.7-2(-3.5) x 0.4-      4) x 0.3-0.5 cm
                           0.8 cm long

Number and shape of    1-2(3) pairs, narrow    1-3 pairs, narrow
lateral lobes of the
leaf

Shape and color of     leaf-like, with 1-3    leaf-like, with 1-3
the bracts                narrow lateral         narrow lateral
                        lobes, greenish or    lobes, green, tipped
                              purple           with red or rarely
                                                     yellow

Length of the                 1-2 mm                5-10 mm
pedicel

Total length of the         1.5-2.2 cm             1.6-3.4 cm
flower

Calyx versus corolla     corolla exserted       corolla exserted
                        from the calyx for     from the calyx for
                              >4 mm                  >4 mm

Color of the calyx      reddish with green    pale green and dark
                       or pale green veins    green to purple tips

Shape and length of     basally inflated,      slightly inflated
the calyx                 12-16(-19) mm        towards its middle
                                              zone, (12-)18-30 mm

Adaxial and abaxial       subequal, the        very unequal, the
clefts of the calyx    adaxial 3-5 mm deep,   adaxial 1-3 mm deep,
                       the abaxial 5-6.5 mm    the abaxial 6-11(-
                               deep               14) mm deep

Lateral clefts of        present, to 1 mm            absent
the calyx                      deep

Color of the corolla    yellow green with        Yellow green,
                         purplish margins       sometimes orange
                                              along the margin of
                                                   the galea

Length and shape of       15-22(-26) mm.      (14-)16-23-(34) mm,
corolla                  slightly curved       slightly incurved
                             outwards

Length of the          8-16 mm, longer than   8-12 mm, longer than
corolla tube                the galea              the galea

Length of the galea          6-10 mm              6-11(-19) mm

Stamens                      included               included

Style and stigma in      style and stigma      exserted from the
anthesis                 included; stigma     corolla and from the
                             capitate            calyx; stigma
                                                 punctiform to
                                                   subclavate

Length of the seeds         1.1-1.3 mm             1.8-2.2 mm

Distribution and          Azuay, Carchi,        Azuay, Bolivar,
altitudinal range          Chimborazo,         Canar, Chimborazo,
                        Cotopaxi, El Oro,      Cotopaxi, El Oro,
                       Imbabura, Pichincha,       Loja, Napo,
                            Tungurahua             Tungurahua
                         (Ecuador); Peru,       (Ecuador); Peru,
                            Bolivia, N        Bolivia. 2500-4000 m
                       Argentina, N Chile.
                           3000-4200 m

(1) Based on Kunth (1817), Holmgren (1984), and the specimens Maguire
& Maguire 61725 (fl, fr), collected in Ecuador, Bolivar, 3900 m, 18
Sep 1969 (COL), and Barclay & Juajibioy 9174, collected in Ecuador,
Napo/Pastaza, 30 Aug 1959 (COL). (2) Based on Edwin (1971) and
Holmgren (1984). (3) Based on Edwin (1971), and Holmgren (1984), and
the holotype (available at http://plants.jstor.org/specimen/
p00587597) and the specimen Barclay 8365 (fl), collected in Ecuador,
Azuay, 3400 m, 30 Jul 1959 (COL).
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Title Annotation:Botanica-taxonomia
Author:Gonzalez, Favio; Pabon-Mora, Natalia
Publication:Caldasia
Date:Jul 1, 2013
Words:4042
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