Additions to the Pantepui Pollen Flora (Venezuelan Guayana): the Maguire Collection

Collectanea Botanica (Barcelona) vol. 29 (2010): 31-49 ISSN: 0010-0730 doi: 10.3989/collectbot.2010.v29.004

Additions to the Pantepui pollen flora (Venezuelan Guayana): the Maguire Collection

C. LÓPEZ-MARTÍNEZ1, A. LARA1, V. RULL1, L. CAMPBELL2 & S. NOGUÉ3

1 Palynology and Paleoecology Lab, Institut Botànic de Barcelona (CSIC-ICUB), Psg. del Migdia s/n., 08038 Barcelona, Spain 2 Plant Research Lab, The New York Botanical Garden, Bronx, NY 10458, USA 3 Long Term Ecology Lab, Oxford University Centre for the Environment, South Parks Road, Oxford OX1 3QY, UK

Author for correspondence: V. Rull ([email protected])

Received 3 September 2010; Accepted 15 November 2010

Abstract This work is a pollen-morphological study of various plant species from Pantepui (Venezuelan Guayana), a region with high biodiversity and endemism, where global warming is expected to have a high impact. The study consists of a series of morphological descriptions of selected taxa from the Maguire pollen reference slide collection of The New York Botanical Garden (NYBG). The collection was initiated under the supervision of Senior Curator Bassett Maguire to advance systematic, palynological, and medical studies; today it has become also useful for other disciplines such as paleoecology, paleoclimatology or forensic studies. The aim of this pollen-morphological study is to enhance the database of pollen descriptions and illustrations for identification purposes, to be used in the ongoing paleoecological reconstructions and, eventually, in other types of studies using pollen, particularly from the Guayanan tepui summits.

Key words: Guayana; Pollen morphology; Taxonomy; Tepuis; Venezuela.

Resumen Adiciones a la flora polínica de Pantepui (Guayana venezolana): la colección Maguire.- Este trabajo es un estudio sobre la morfología polínica de varias especies de plantas de Pantepui (Guayana venezolana), una región con elevada biodiversidad y un alto grado de endemismo, donde se espera un fuerte impacto del calentamiento global. El estudio consiste en una serie de descripciones morfológicas de taxones pertenecientes a la colección Maguire de láminas de referencia del Jardín Botánico de Nueva York (NYBG). La colección fue iniciada bajo la dirección del Curador Jefe Basset Maguire con el objeto de avanzar en estudios de sistemática, palinología y medicina; aunque actualmente también resulta útil en otras disciplinas, como por ejemplo la paleoecología, la paleoclimatología o los estudios forenses. El objetivo de este estudio de morfología polínica es aumentar la base de datos de descripciones e ilustraciones polínicas con propósitos de identificación taxonómica, para ser utilizada en reconstrucciones paleoecológicas y otro tipode estudios actualmente en desarrollo, en particular en las cimas de los tepuis de Guayana.

Palabras clave: Guayana; Morfología polínica; Taxonomía; Tepuis; Venezuela. 32 C. LÓPEZ-MARTÍNEZ, A. LARA, V. RULL, L. CAMPBELL & S. NOGUÉ

INTRODUCTION and with a surface of < 1 to > 1000 km2 (typically 200-500 km2) (Huber, 1995a). The tepui summits are This paper is part of a series devoted to the pollen- more or less flat and hold unique and specialized life morphological study of vascular plant species from forms and communities, as well as a high diversity the Venezuelan Guayana (Fig. 1), particularly from and an amazing degree of endemism (Rull, 2009). the characteristic sandstone tabular mountains This biotic singularity led to the creation of the Pan- (tepuis), initiated by Salgado-Labouriau & Villar tepui biogeographical province, which embraces all (1992) and continued by Rull (2003). The biodi- the tepui summits above 1500 m elevation (Huber, versity and endemism of this region are striking, 1988, 1994). Vascular plants are the better known but the origin and evolution of such exceptional organisms of Pantepui, with almost 2500 species, biotic features has been object of intense debate, of which > 60% are endemic to the Guayana region which is still ongoing (Rull, 2010). On the other and > 40% are endemic to Pantepui, including 25% hand, the tepuian flora is expected to be seriously species endemic to a single tepui (Berry & Riina, affected by relatively rapid biodiversity loss under 2005). Similarly, most tepuian vegetation types are ongoing global warming. Hence, paleoecological unparalleled worldwide (Huber, 2005). Pantepui and paleoclimatic studies on this region can provide is mostly pristine due to its remoteness and inac- useful information about biogeographical and evo- cessibility; only a few of the tepui summits can be lutionary processes as well as the effects of climate reached by foot after several days of walking and change on the tepuian biota. climbing. Indigenous people living in the surroun- Besides the papers mentioned above, which are ding lowlands and midlands do not visit the tepui monographic for the pollen of the Guayana region, summits because they consider them the home of the available literature dealing with this subject is gods and access is not allowed. On the other hand, scarce and organized taxonomically. Rapateaceae there is no potential for commercial profit through and Asteraceae are the most thoroughly studied activities such as mining, agriculture, cattle raising, families in this sense, although the knowledge of As- etc. (Gorzula & Huber, 1992; Huber, 1995b). Tourism teraceae is restricted to some genera of Mutisioideae and biopiracy are restricted by severe regulations that, (Carlquist, 1961; Tellería, 2008; Funk et al., 2009; unfortunately, also hinder scientific research (Rull Ubiergo et al., 2009). Accordingly, the aim of this & Vegas-Vilarrúbia, 2008; Rull et al., 2008a). The study, as well as the previously mentioned works, main threat for the Pantepui biota is global warming is to enhance the database of pollen descriptions that, according to recent estimates, would determine and illustrations for identification purposes, to be significant biodiversity losses by upward migration used in the ongoing pollen-based paleoecological and the corresponding habitat loss and fragmentation reconstructions (Rull et al., 2010) and, eventually, for many endemic species (Rull & Vegas-Vilarrúbia, in other types of studies using pollen. This work 2006; Rull et al., 2008b; Nogué et al., 2009). is based on the Maguire pollen reference slide collection, initiated in the forties of the past century by Senior Curator B. Maguire of the New York MATERIAL AND METHODS Botanical Garden (NYBG) to make progress in biological research. Most genera and/or species Slides for this study are part of the Maguire pollen studied here are endemic to the Guayana region reference slide collection preserved in The New York or even to the Pantepui province or a single tepui. Botanical Garden (NYBG; see http://www.nybg.org/). The collection comprises 3650 preparations of mod- Study area: Pantepui ern pollen representing 153 vascular plant families, of which 11 are non-seed plants. Some preparations In the local indigenous language (Pemón), “tepui” include duplicate slides. Most vouchers are deposited means “stone bud”, and is the name for the cha- in the Steere Herbarium of NYBG and can be viewed racteristic sandstone/quartzite tabular mountains on-line (http://www.nybg.org/). The collection was of Guayana (Fig. 2). They vary in altitude and initiated under the supervision of Senior Curator extension, ranging from around 1100 to 3000 m Bassett Maguire (1904–1991) to advance systematic, (typically 2000-2600 m) of maximum elevation, palynological, and medical studies. Research utilizing

Collectanea Botanica vol. 29 (2010): 31-49, ISSN: 0010-0730, doi: 10.3989/collectbot.2010.v29.004 Tepuian pollen flora (Venezuelan Guayana) 33

10º 500 km VENEZUELA Orinoco

GUYANA

SURINAME COLOMBIA FRENCH GUIANA

0º ECUADOR

Amazon PERU BRAZIL

80º 70º 60º 50º Figure 1. Map of northern South America, showing the approximate extension of the Guayana Shield (in grey) and the Precambrian sandstones and quartzites of the Roraima group (in black), on which the tepuis are modelled (Briceño & Schubert, 1990). The region under study (the Venezuelan Guayana) is indicated by a box.

Figure 2. Example of a tepui: picture of the Roraima tepui [Photograph by V. Rull].

Collectanea Botanica vol. 29 (2010): 31-49, ISSN: 0010-0730, doi: 10.3989/collectbot.2010.v29.004 34 C. LÓPEZ-MARTÍNEZ, A. LARA, V. RULL, L. CAMPBELL & S. NOGUÉ these specimens will undoubtedly surpass the original genera from the Venezuelan Guayana are described, objectives by serving fields as for example paleoecol- if any. The following abbreviations have been used ogy, climate change or forensic studies, among others. in the text and descriptions: PV, polar view; EV, The breadth of the collection is international in scope equatorial view; P/E, ratio of the length of the polar with the greatest geographic representation in tropical axis (P) to the equatorial diameter (E) (this ratio is America, particularly the Guayana region of northern the basis of a system of shape classes, Punt et al., South America. Systematic strengths of the collec- 2007). Finally, the photographs are ordered by pollen tion reflect Maguire’s interest in plant groups with morphology (monads, tetrads and polyads, and the high species diversity or endemism in the Guayana number and type of apertures). The scale bar in the (e.g., Clusiaceae, Gentianaceae, Rapateaceae, and plates is always 10 mm long, except for the images Velloziaceae). Samples for the Maguire collection of Moronobea jenmanii Engl. in Mart. var. fanshawei were mostly obtained from dried or fluid preserved Maguire and Irlbachia cardonae (Gleason) Maguire, collections housed at NYBG, but also include dried where it is 20 mm long. or fresh material as well as prepared slides from other institutions. Pollen grains were chemically treated with POLLEN DESCRIPTIONS KOH, acetolysis, a combination of the two, or were untreated, and mounted in glycerin jelly. The slides Asteraceae are stored in vertical sleeves in a steel cabinet (see Howard & Boom, 1990, Fig. 11). Glossarion rhodanthum Maguire & Wurdack Measurements for this work have been taken in (Plate 1: 6-9) all the available well-preserved grains; the number (NY 2584; B. Maguire et al. 37126; Neblina) of grains used in each case is indicated after the Monads. Isopolar, subprolate (P/E = 1.27), amb measurements (n=x). Measurements were taken using circular slightly lobate. Tricolporate, zonoaperturate. the photographic software DPXViewer 1.14.9 and Ectocolpi long, sharp ends, fusiform (74.2 x 4.2 mm), a DeltaPix InfinityX camera. A number of species marginate. Ora fusiform, lalongate (7.4 x 25.8 mm). could not have been described because of the absence Exine 8.8 mm thick (excluding spines), crescentic of suitable pollen grains due to bad preservation. (in polar view, very thick exine in the medium of These taxa are listed in the appendix for reference. the intercolpium, gradually thinning towards the Taxa are ordered alphabetically by families, follow- colpi), caveate, tectate, echinate, supratectal spines ing the classification of the Flora of the Venezuelan triangular (ca. 3 mm long and 3 mm wide at the base) Guayana (Steyermark et al., 1995-2005). For each with slightly rounded ends. Collumellae gradually plant species there is a morphological description more dense and/or smaller near the margo. Sexine accompanied by some photographs of the pollen thicker than nexine. Grain size (excluding spines): grains found in the slide. The descriptions follow 89.8-102.8 x 71.4-80.5 mm, n = 10. the Rull (2003) style for consistency, and use the standard nomenclature system of Punt et al. (2007). Gongylolepis benthamiana R. Schomb. (Plate 1: The title of every description summarizes the genus 10-13) and species, the species authors, the pollen cata- (NY 2586; T. Koyama & G. Agostini 7366; logue number, the collector, the collection location, Venezuela) as well as other information present on the label Monads. Isopolar, subprolate to prolate of the slide. Within the description, colpi and ora (P/E = 1.33), amb circular slightly lobate. Tricol- sizes are expressed as length x width (regarding the porate, zonoaperturate. Ectocolpi long, rounded direction of the polar axis, in equatorial view), and ends, elliptic (83.2 x 4.2 mm), margo thick (4.0 mm), grain size, as length of the polar axis x equatorial psilate. Ora elliptic, lolongate (19.9 x 12.3 mm). diameter. Also, the more recently accepted name of Exine 9.3 mm thick (excluding spines), caveate, the species is noted where necessary, as well as the tectate, echinate, supratectal spines triangular (ca. literature where pollen of other species of the same 4 mm long and ca. 5 mm at the base) with slightly

Plate 1. 1-3, Kunhardtia rhodantha (Rapateaceae); 4-5, Barbacenia magalhaesii (Velloziaceae); 6-9, Glossarion rhodanthum (Asteraceae); 10-13 Gongylolepis benthamiana (Asteraceae).

Collectanea Botanica vol. 29 (2010): 31-49, ISSN: 0010-0730, doi: 10.3989/collectbot.2010.v29.004 Tepuian pollen flora (Venezuelan Guayana) 35

3 4 5

1 2

6 7

10 mm 9

10 11

12 13

Collectanea Botanica vol. 29 (2010): 31-49, ISSN: 0010-0730, doi: 10.3989/collectbot.2010.v29.004 36 C. LÓPEZ-MARTÍNEZ, A. LARA, V. RULL, L. CAMPBELL & S. NOGUÉ rounded ends. Sexine thicker than nexine. Grain Clethraceae size: 112.4-133.6 x 87.2-103.0 mm (excluding spines), n = 11. Clethra alnifolia L. (Plate 2: 25-28) (NY 3008; Martha’s Vineyard, Mass, 18 Sept Stenopadus cucullatus Maguire (Plate 2: 14-17) 1964; F. C. Mackoever) (NY 2592; B. Maguire & C. K. Maguire 35029; Monads. Isopolar, prolate-spheroidal (P/E = 1.13), Venezuela) amb circular. Tricolporate, zonoaperturate. Ectocolpi Monads. Isopolar, prolate (P/E = 1.53), amb circu- long (21.6 mm), shape difficult to distinguish. Ora lar slightly lobate. Tricolporate, zonoaperturate. Ec- lalongate, constricted, fastigium present. Exine tocolpi long, rounded ends, elliptic (74.8 x 3.7 mm), 2.0 mm thick, tectate, psilate. Sexine and nexine slightly marginate. Ora fusiform, lalongate approximately of the same thickness. Grain size: (7.6 x 23.6 mm). Exine 3.1 mm thick, thicker at the 25.9-32.9 x 23.4-30.1 mm, n = 10. poles (4.8 mm), tectate, psilate (columellae visible Comments: Clethra guianensis Klotzsch ex. through the tectum). Sexine thicker than nexine. Meisn. is described in Rull (2003). Grain size: 91.5-101.5 x 56.6-69.9 mm, n = 11. Comments: Stenopadus sp. described in Rull Clusiaceae (2003). Clusia cochlitheca Maguire (Plate 2: 29-30) (NY 3476; B. Maguire 36900) Bonnetiaceae Monads. Amb triangular, angulaperturate. Tricol- Bonnetia celiae Maguire (Plate 2: 18-21) porate, operculate (operculum not always present), (NY 957; B. Maguire 35327; Venezuela) zonoaperturate. Ectocolpi short (12.3 mm), ora diffi- Monads. Isopolar, subprolate (P/E = 1.31), cult to see. Exine 1.1 mm thick, semitectate, perforate. amb sub-triangular, angulaperturate. Tricolporate, Sexine and nexine indistinguishable for measuring. zonoaperturate. Ectocolpi long, sharp ends, fusiform Grain size: 28.3-32.7 mm, median in PV, n = 5. (50.8 x 9.7 mm), marginate. Ora elliptic, lolongate Comments: Description based on grains in PV. (12.8 x 8.8 mm), annulus present. Exine 4.7 mm Clusia melchiorii Gleason is described in Rull (2003). thick, semitectate, perforate. Sexine thicker than nexine. Grain size: 54.0-74.4 x 43.1-57.3 mm, n = 9. Clusia orthoneura Standl. (Plate 2: 31-34) Comments: Bonnetia lanceifolia Maguire, B. (NY 3483; B. Maguire 62205) roraimae Oliver and B. sessilis Benth. are des- Monads. Isopolar, subprolate (P/E = 1.25), amb cribed in Salgado-Labouriau & Villar (1992) and triangular-circular, angulaperturate. Tricolporate, Rull (2003). operculate (operculum not always present), zonoaperturate. Ectocolpi long (29.5 mm), marginate, Bonnetia roraimae Oliv. ex Thurn (Plate 2: 22-24) constricted in the equatorial region, sharp ends. (104; Maguire 33421) Ora difficult to distinguish. Exine 1.8 mm thick, Monads. Isopolar, oblate-spheroidal (P/E = 0.97), semitectate, reticulate, heterobrochate (lumina amb circular-triangular, angulaperturate. Tricolpo- slightly smaller near the colpi). Sexine slightly rate, zonoaperturate. Ectocolpi long, sharp ends, thiner than nexine. Grain size: 31.5-37.6 x 25.3- fusiform (27.4 mm x 3.8 mm). Ora circular (9.3 mm 32.4 mm, n = 20. diameter), sometimes operculate. Exine 2 mm thick, semitectate, perforate. Sexine thicker than nexine. Hypericum uliginosum H. B. K. (Plate 2: 35-36) Grain size: 29.1-41.2 x 27.3-43.0 mm, n = 6. (NY 753; E. P. Killip & A. C. Smith 20487; Comments: Bonnetia roraimae is described in Colombia) Salgado-Labouriau & Villar (1992) and Rull (2003). Monads. Isopolar, subprolate (P/E = 1.22).

Plate 2. 14-17, Stenopadus cucullatus (Asteraceae); 18-21, Bonnetia celiae (Bonnetiaceae); 22-24, Bonnetia roraimae (Bonnetiaceae); 25-28, Clethra alnifolia (Clethraceae); 29-30, Clusia cochlitheca (Clusiaceae); 31-34 Clusia orthoneura (Clusiaceae); 35-36, Hypericum uliginosum (Clusiaceae); 37-40, Neotatea neblinae (Clusiaceae); 41-43, Tovomita weddelliana (Clusiaceae); 44-46, Tapeinostemon longiflorum (Gentianaceae).

Collectanea Botanica vol. 29 (2010): 31-49, ISSN: 0010-0730, doi: 10.3989/collectbot.2010.v29.004 Tepuian pollen flora (Venezuelan Guayana) 37

18 19

14 15

20 21

16 17 22

25 26 27 28 29 30

31 32 33 34

35 42 41

37 38 10 mm

43 36

39 40 44 45 46

Collectanea Botanica vol. 29 (2010): 31-49, ISSN: 0010-0730, doi: 10.3989/collectbot.2010.v29.004 38 C. LÓPEZ-MARTÍNEZ, A. LARA, V. RULL, L. CAMPBELL & S. NOGUÉ

Tricolporate, zonoaperturate. Ectocolpi long (27.5 Droseraceae mm), rounded ends, marginate, ora difficult to see. Exine 2.3 mm thick, semitectate, perforate. Sexine Drosera roraimae (Klotzsch ex Diels) Maguire thicker than nexine. Grain size: 46.5-51.4 x 38.5- (Plate 4: 89-91) 41.6 mm, n = 4. (NY 2781; B. Maguire 42338) Tetrads, uniplanar (rhomboidal and tetragonal) or Moronobea jenmanii Engl. in Mart. var. fanshawei multiplanar (decussate and tetrahedral). Individual Maguire (Plate 3: 82-84) grains inaperturate, heteropolar (distal pole clavate- (NY 2315; S. S. Tillett & C. L. Tillett 44904; B. microechinate and proximal pole conspicuously Guiana) folded), oblate-suboblate (P/E = 0.75), amb subcir- Monads. Isopolar, subprolate (P/E = 1.18), cular. Exine 3.5 mm thick, intectate, microechinate amb circular-quadrangular angulaperturate. Te- and clavate. Sexine thicker than nexine. Individual tracolporate (sometimes pentacolporate), zonoa- grain size: 33.0-44.3 x 46.6-54.2 mm, n = 16. Tetrad perturate. Ectocolpi 94.3 mm long and narrow. diameter: 60.3-74.4 mm, n = 8. Ora difficult to see. Exine 4.7 mm thick, psilate, Comments: Drosera sp. described in Salgado- tectate. Sexine and nexine approximately of the Labouriau & Villar (1992) and Rull (2003). same thickness. Grain size: 169.2-184.0 x 140.0- 166.3 mm, n = 10. Ericaceae

Neotatea neblinae Maguire (Plate 2: 37-40) Bejaria glauca Humb. & Bonpl. var. coarctata (NY 2415; B. Maguire 42178; Neblina) Mansf. & Sleumer (Plate 4: 92-94) Monads. Isopolar, prolate-spheroidal (P/E = 1.04), (NY 1391; B. Maguire & C. K. Maguire # 44345; amb triangular, angulaperturate. Tricolporate, Ecuador) zonoaperturate. Ectocolpi long, fusiform, sharp ends Tetrads, multiplanar, tetrahedral or decussate, out- (47.8 x 4.3 mm). Ora lalongate (15.1 x 28.3 mm), line circular slightly lobate. Viscin threads absent. slightly constricted in the middle, fastigium present. Individual grains heteropolar, oblate (P/E = 0.74), Exine 3.3 mm thick, semitectate, perforate. Sexine amb circular, tricolporate. Half ectocolpi 8.1 mm slightly thicker than nexine. Grain size: 54.2- long, narrow, slightly marginate. Ora difficult to dis- 67.2 x 53.3-65.3 mm, n = 10. tinguish. Exine 2.3 mm thick, psilate, tectate. Sexine and nexine approximately of the same thickness. Tovomita weddelliana Planch. & Triana (Plate 2: Individual grain size: 30.5-37.3 x 41.1-48.8 mm, 41-43) n = 10. Tetrad diameter: 56.0-67.3 mm, n = 10. (NY 3528; L. Aristeguieta & G. Agostini 6344; Comments: Bejaria aestuans Mutis ex L., in Venezuela) Steyermark et al. (1995-2005). Monads. Isopolar, prolate (P/E = 1.47), amb circular lobate, fossaperturate. Tricolporate, zo- Cavendishia duidae A. C. Sm. in Gleason (Plate 5: noaperturate. Ectocolpi long (28.0 mm), marginate, 104-105) constricted in the equatorial region, sharp ends. (NY 1399; Maguire et al. 29707; Venezuela) Ora only visible in optical section. Exine 1.7 mm Tetrads, multiplanar tetrahedral or decussate, thick, semitectate, perforate. Sexine and nexine outline circular-triangular slightly lobate. Viscin approximately of the same thickness. Grain size: threads absent. Individual grains heteropolar, oblate 29.9-37.1 x 20.5-25.6 mm, n = 10. (P/E = 0.72), amb circular, tricolporate. Half ectocolpi

Plate 3. 47-50, Chorisepalum ovatum var. ovatum (Gentianaceae); 51-53, Macrocarpaea neblinae (Gentianaceae); 54- 55, Psittacanthus obovatus (Loranthaceae); 56-58, Diacidia galphimioides (Malpighiaceae); 59-60, Myrcia bolivarenses (Myrtaceae); 61-62, Ouratea gillyana (Ochnaceae); 63-64, Freziera sericea (Ternstroemiaceae); 65-66, Philacra duidae (Ochnaceae); 67-69, Philacra steyermarkii (Ochnaceae); 70, Poecilandra pumila (Ochnaceae); 71, Tyleria spathulata (Ochnaceae); 72, Schoepfia flexuosa (Olacaceae); 73-74, Duranta mutisii (Verbenaceae); 75-76, Banisteriopsis inebrians (Malpighiaceae); 77-79, Borreria laevis (Rubiaceae); 80-81, Heliamphora neblinae var. neblinae (Sarraceniaceae); 82-84, Moronobea jenmanii var. fanshawei (Clusiaceae).

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50 47 48 49 57

51 53 54 55

65 63

61 62 64 66 59 60 52

67 68 69 70 71 72 77

78 74 75 76 73

10 mm

81 80 79

20 mm

83 84

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9.5 mm long, narrow, slightly marginate. Ora difficult ine and nexine approximately of the same thickness. to distinguish. Exine 1.6 mm thick, psilate, tectate. Individual grain size: 39.3-48.4 x 43.6-53.6 mm, n = Sexine and nexine indistinguishable for measuring. 10. Tetrad diameter: 71.6-80.5 mm, n = 7. Individual grain size: 32.2-38.2 x 40.8-55.4 mm, n=10. Tetrad diameter: 57.8-71.8 mm, n = 10. Notopora schomburgkii Hook.f. (Plate 6: 115-116) (NY 1668; B. Maguire et al. 46150A; British Disterigma humboldtii Nied. (Plate 5: 108-110) Guiana) (NY 1406; J. A. Steyermark, J. J. Wurdack Tetrads, multiplanar, tetrahedral or decussate, out- 1156; Venezuela) line triangular or quadrangular. Viscin threads absent. Tetrads, multiplanar, tetrahedral or decussate, Individual grains heteropolar, oblate (P/E = 0.74), outline circular-triangular. Viscin threads absent. In- amb circular, tricolporate. Half ectocolpi 20.3 mm dividual grains heteropolar, suboblate (P/E = 0.80), long, narrow, marginate. Ora difficult to distinguish. amb circular, tricolporate. Half ectocolpi 17.4 mm Exine 3.1 mm thick, psilate, tectate. Sexine and nex- long, narrow, marginate. Ora difficult to distinguish. ine approximately of the same thickness. Individual Exine 2.7 mm thick, psilate, tectate. Sexine and grain size: 40.5-49.9 x 56.5-63.7 mm, n = 10. Tetrad nexine indistinguishable for measuring. Individual diameter: 75.7-90.4 mm, n = 10. grain size: 31.1-35.6 x 37.4-47.5 mm, n = 19. Tetrad diameter: 51.5-61.4 mm, n = 19. Notopora smithiana Steyerm. & Maguire (Plate 6: 117-120) Ledothamnus atroadenus Maguire, Steyerm. & (NY 3494; J. A. Steyermark & J. J. Wurdack Luteyn var. atroadenus (Plate 5: 111-112) 823; Chimantá, Venezuela) (NY 1413; J. J. Wurdack # 34190; Venezuela) Tetrads, multiplanar, tetrahedral or decussate, Tetrads, multiplanar, tetrahedral or decussate, out- outline circular-triangular. Viscin threads absent. line circular slightly lobate (sometimes triangular- Individual grains heteropolar, oblate (P/E = 0.72), circular). Viscin threads absent. Individual grains amb circular, tricolporate. Half ectocolpi 19.1 mm heteropolar, oblate (P/E = 0.73), amb circular, long, 3.8 mm wide, sharp ends, sometimes appear tricolporate. Half ectocolpi 10.8 mm long, narrow, narrow. Ora difficult to distinguish, probably marginate. Ora difficult to distinguish. Exine 2.8m m lalongate (10.4 mm wide). Exine 2.9 mm thick, thick, psilate, tectate. Sexine and nexine approxi- microrugulate, tectate. Sexine and nexine approximately of the same thickness. Individual grain size: mately of the same thickness. Individual grain size: 28.8-34.6 x 40.2-45.8 mm, n=11. Tetrad diameter: 37.3-42.1 x 52.5-59.0 mm, n = 11. Tetrad diameter: 51.3-60.7 mm, n = 11. 70.1-77.3 mm, n = 11. Comments: Ledothamnus luteus Maguire, Steyerm. & Luteyn described in Salgado-Labouriau Orthaea paruensis Spruce (Plate 6: 121-124) & Villar (1992) and Rull (2003). (NY 1424; K. D. Phelps & C. B. Hitchcock 477; Venezuela) Mycerinus chimantensis Maguire, Steyerm. & Tetrads, multiplanar, tetrahedral or decussate, Luteyn (Plate 5: 106-107) outline circular-triangular. Viscin threads absent. (NY 1422; J. A. Steyermark & J. J. Wurdack Individual grains heteropolar, suboblate (P/E = 0.77), 825; Venezuela) amb circular, tricolporate. Half ectocolpi 13.6 mm Tetrads, multiplanar, tetrahedral or decussate, long, narrow, marginate (half margo: 16.2 x 3.3 mm). outline circular slightly lobate. Viscin threads absent. Ora difficult to distinguish but they seem lalongate Individual grains heteropolar, suboblate (P/E = 0.84), (10.7 mm wide). Exine 2.7 mm thick, psilate, tectate. amb circular, tricolporate. Half ectocolpi 16.8 mm Sexine and nexine approximately of the same thick- long but often difficult to distinguish. Ora difficult to ness. Individual grain size: 30.1-34.7 x 39.1-45.7 mm, distinguish. Exine 2.2 mm thick, psilate, tectate. Sex- n = 12. Tetrad diameter: 55.9-62.1 mm, n = 12.

Plate 4. 85-86, Utricularia foliosa (Lentibulariaceae); 87-88, Monnina chanduyensis (Polygalaceae); 89-91, Drosera roraimae (Droseraceae); 92-94, Befaria glauca var. coarctata (Ericaceae); 95, Symbolanthus elisabethae (Gentianaceae); 96-99, Schultesia heterophylla (Gentianaceae).

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85 86 90

87 88 89

92 93

10 mm

95 98 99

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Tepuia cardonae A. C. Sm. (Plate 5: 113-114) Irlbachia cardonae (Gleason) Maguire (Plate 6: (NY 1671; J. A. Steyermark & J. J. Wurdack 127-129) 1039; Venezuela) (NY 2531; B. Maguire et al. 33917; Venezuela) Tetrads, multiplanar, tetrahedral or decussate, out- Polyad, constituted by 16 grains, outline circular. line circular or subtriangular. Viscin threads absent. Individual grains heteropolar (only distal pole echi- Individual grains heteropolar, oblate (P/E = 0.69), nate), suboblate (P/E = 0.78), amb circular. Apertures amb circular, tricolporate. Half ectocolpi 10.8 mm difficult to distinguish due to the sculpture (Gentiana- long, narrow. Ora difficult to distinguish. Exine ceae are normally tricolporate). Exine 4.7 mm thick 3.2 mm thick, psilate, tectate. Sexine and nexine (excluding spines), reticulate, semitectate, echinate approximately of the same thickness. Individual (4-5 big curved spines per grain with rounded ends: grain size: 24.1-39.1 x 38.5-54.9 mm, n = 9. Tetrad 22.5 mm long, 13.4 mm base diameter), supratectal diameter: 47.6-62.6 mm, n = 9. granula and verrucae irregularly distributed. Sexine Comments: Tepuia venusta Camp. described thicker than nexine. Individual grain size: 43.0-57.8 in Salgado-Labouriau & Villar (1992) and Rull x 56.9-69.7 mm, n = 7. Polyad diameter 190.9-210.2 (2003). mm (excluding spines), n = 5.

Thibaudia cupatensis Huber (Plate 6: 125-126) Macrocarpaea neblinae Maguire & Steyerm. (NY 1436; B. Maguire et al. # 42263; Venezuela) (Plate 3: 51-53) Tetrads, multiplanar, tetrahedral or decussate, out- NY 2723; B. Maguire et al. 37103; Venezuela) line circular, slightly lobate. Viscin threads absent. Monads. Isopolar, prolate-spheroidal (P/E = 1.06), Individual grains heteropolar, suboblate (P/E = 0.76), amb circular. Tricolporate, zonoaperturate. Ectocolpi amb circular, tricolporate. Half ectocolpi 16.8 mm long, sharp ends, sometimes slightly rounded, long, narrow, marginate. Ora difficult to distinguish. fusiform (28.5 x 2.3 mm), marginate. Ora elliptic Exine 2.4 mm thick, psilate, tectate. Sexine and nex- (6.5 x 4.3 mm), lolongate. Exine 3.8 mm thick, semi- ine approximately of the same thickness. Individual tectate, reticulate irregularly heterobrochate, simpli- grain size: 39.4-46.9 x 52.3-58.6 mm, n = 10. Tetrad and duplicolumellate. Sexine thicker than nexine. diameter: 68.2-78.8 mm, n = 10. Grain size: 41.7-45.1 x 38.8-43.0 mm, n = 10. Comments: Thibaudia nutans Klotzsch ex Mansf. Comments: Macrocarpaea tepuiensis (Gleason) described in Salgado-Labouriau & Villar (1992) Steyermark described in Rull (2003). and Rull (2003). Neblinantha neblinae Maguire (Plate 5: 100-103) (NY 2541; B. Maguire et al. 60527; Neblina) Gentianaceae Tetrads, multiplanar, tetrahedral or decussate. In- dividual grains heteropolar, oblate (P/E = 0.72), amb Chorisepalum ovatum Gleason var. ovatum circular-slightly lobate. Tricolporate, colpi and ora (Plate 3: 47-50) difficult to observe due to the sculpture pattern. Exine (NY 2500; B. Maguire et al. 27921; Venezuela) 7.7 mm thick, semitectate, reticulate, heterobrochate, Monads. Isopolar, subprolate (P/E = 1.21), amb simplicolumellate. Sexine thicker than nexine. Indi- circular. Tricolporate, zonoaperturate. Ectocolpi vidual grain size: 47.5-54.0 x 63.2-77.9 mm, n = 10. long and wide (36.1 x 3.9 mm), sharp ends, cons- Tetrad diameter: 90.3-103.8 mm, n = 10. tricted at the equatorial region. Ora apparently lolongate, but difficult to distinguish. Exine 4.5 Schultesia heterophylla Miq. (Plate 4: 96-99) mm thick, semitectate, reticulate, irregularly he- (NY 2565; J. J. Wurdack & N. G. L. Guppy 15; terobrochate (lumina smaller in polar regions), Venezuela) simplicolumellate. Sexine thicker than nexine. Tetrads, multiplanar, tetrahedral or decussate, Grain size: 48.0-54.2 x 35.0-44.9 mm, n = 6. outline circular-triangular. Individual grains hetero-

Plate 5. 100-103, Neblinantha neblinae (Gentianaceae); 104-105, Cavendishia duidae (Ericaceae); 106-107, Mycerinus chimantensis (Ericaceae); 108-110, Disterigma humboldtii (Ericaceae); 111-112, Ledothamnus atroadenus var. atroadenus (Ericaceae); 113-114, Tepuia cardonae (Ericaceae).

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103 104

100

106 105

101

108

107

110

10 mm 109 102

111 112 113 114

Collectanea Botanica vol. 29 (2010): 31-49, ISSN: 0010-0730, doi: 10.3989/collectbot.2010.v29.004 44 C. LÓPEZ-MARTÍNEZ, A. LARA, V. RULL, L. CAMPBELL & S. NOGUÉ polar, oblate to suboblate (P/E = 0.75), amb circular. turate. Tricolp(or)ate, zonoaperturate, syncolp(or) Triporate, pori elliptic (7.2 mm long, 8.9 mm wide), ate. Exine psilate, tectate. annulate. Exine 8.0 mm thick, semitectate, reticulate Comments: One single pollen grain found (PV). heterobrochate. Sexine thicker than nexine. Indivi- dual grain size: 46.7-54.0 x 63.4-73.3 mm, n = 11. Tetrad diameter: 88.3-97.9 mm, n = 11. Malpighiaceae Banisteriopsis inebrians C. V. Morton (Plate 3: Symbolanthus elisabethae Gilg (Plate 4: 95) 75-76) (NY 1266; J. A. Steyermark 75662; Venezuela, (NY 2703; B. A. Krukoff-8550; Brazil) Bolivar, June 1953) Monads. Spheroidal. Pantoporate, with ir- Tetrads, multiplanar, tetrahedral or decussate. Ap- regular colpoids in several directions. Pori elliptic ertures difficult to observe due to the sculptural (3.4 x 2.1 mm), with diffuse outline. Exine 3.8 mm pattern. Exine 5-6 mm thick, semitectate, reticulate thick, tectate, psilate. Sexine thicker than nexine. heterobrochate, simplicolumellate. Sexine thicker Grain diameter: 37.5-46.2 mm, n = 10. than nexine. Tetrad diameter: 85.8-90.0 mm, n = 4. Diacidia galphimioides Griseb (Plate 3: 56-58) Tapeinostemon longiflorumMaguire & Steyerm. (NY 670; B. Maguire 44091) (Plate 2: 44-46) Monads. Isopolar, prolate-spheroidal (P/E = 1.04), (NY 2579; J. A. Steyermark 103937; Venezuela) amb circular-triangular, angulaperturate. Tricolpo- Monads. Isopolar, subprolate (P/E = 1.30), amb rate, zonoaperturate. Ectocolpi short, sharp ends, circular. Tricolporate, zonoaperturate. Ectocolpi fusiform (13.2 x 3.2 mm). Ora elliptic, lalongate long (30.3 mm) and narrow, sharp ends. Ora circular (4.4 x 6.2 mm), fastigium present. Exine 1.7 mm (6.4 mm diameter). Exine 3.0 mm thick, semitectate, thick, semitectate, reticulate heterobrochate, lumina perforate. Sexine thicker than nexine. Grain size: size decreasing towards the apertures and the polar 43.3-53.9 x 35.6-42.0 mm, n = 6. region. Nexine slightly thicker than sexine. Grain size: 24.7-28.9 x 22.9-27.4 mm, n = 11. Lentibulariaceae Comments: The slide has the epithet duckei, however this name was never published. Utricularia foliosa L. (Plate 4: 85-86) (NY 1646; E. Asplund 5793; Ecuador) Monads. Isopolar, subprolate (P/E = 1.29), amb Myrtaceae circular slightly multilobate. Polyzonocolporate Myrcia bolivarensis (Steyerm.) McVaugh (Plate (17 apertures). Ectocolpi long (68.7 mm) and narrow, 3: 59-60) endocingulum present (zonorate). Exine 2.0 mm (NY 2193; S. S. Tillett & C. L. Tillett 45012; B. thick, psilate. Sexine and nexine indistinguishable. Guiana) Grain size: 81.63-88.1 x 63.9-66.5 mm, n = 3. Monads. Isopolar, amb triangular, angulaper- Comments: Utricularia quelchii N. E. Brown turate. Tricolporate, syncolporate. Exine 1.7 mm described in Rull (2003) and Salgado-Labouriau thick, reticulate or areolate with ondulating surface & Villar (1992). (difficult to observe due to bad preservation). Sexine and nexine difficult to distinguish for measuring. Loranthaceae Grain size: 29.2-33.8 mm, median in PV, n = 10. Comments: Description based on grains in PV. Psittacanthus obovatus Eichl. (Plate 3: 54-55) Myrcia bonnetia-sylvestris (Steyerm.) Steyermark (NY 1612; P. C. Hutchinson 1363 H. G. H.; Peru) described in Salgado-Labouriau & Villar (1992) Monads. Amb subtriangular concave, angulaper- and Rull (2003).

Plate 6. 115-116, Notopora schomburgkii (Ericaceae); 117-120, Notopora smithiana (Ericaceae); 121-124, Orthaea paruensis (Ericaceae); 125-126, Thibaudia cupatensis (Ericaceae); 127-129, Irlbachia cardonae (Gentianaceae).

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117 115 116

120 118 119

121 122 123 124

10 mm

125 127

128 129

20 mm 126

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Ochnaceae mm, n = 1), sharp ends, slightly marginate. Ora elliptic, lalongate (3.2 x 6.4, n = 1). Exine 1.3 mm thick, psilate, Ouratea gillyana (Dwyer) Sandwith & Maguire tectate. Sexine and nexine approximately of the same (Plate 3: 61-62) thickness. Grain size: 24.0-32.5 x 20.2-26.4 mm, n = 11. (B. Maguire & D. B. Fanshawe 32629; 311) Monads. Isopolar, spheroidal (P/E = 1.00), amb circular. Tricolporate, zonoaperturate. Ecto- Olacaceae colpi apparently fusiform but difficult to observe Schoepfia flexuosa Roem. & Schult. (Plate 3: 72) (9.9 mm long, n=2), ora slightly elliptic lolongate (NY 3037; J. A. Steyermark & J. J. Wurdack (4.7 x 3.9 mm). Exine 1.4 mm thick, psilate and tec- 326, Feb. 2, 1955; Chimantá, Venezuela) tate (columellae visible through the tectum). Sexine Monads. Isopolar, amb triangular-rounded, an- and nexine approximately of the same thickness. gulaperturate. Tricolporate, syncolporate. Exine Grain size: 23.2-26.7 x 23.1-27.1 mm, n = 10. 1.52 mm thick (measured in PV), psilate. Sexine Philacra duidae (Gleason) Dwyer (Plate 3: 65-66) and nexine approximately of the same thickness. (NY 279; G. H. H. Tate 529) Grain size: 25.7-32.1 mm, median in PV, n = 6. Monads. Isopolar, prolate (P/E = 1.36), amb circular. Comments: Described in Maguire et al. (1974), Tricolporate, zonoaperturate. Ectocolpi long (19.5 mm) on the same slide. and narrow. Ora elliptic, lalongate: 3.5 mm long (n = 7) x 7.9 mm wide (n = 4). Exine 1.0 mm thick, psilate Polygalaceae and tectate (columellae visible through the tectum). Sexine and nexine indistinguishable for measuring. Monnina chanduyensis Chod. (Plate 4: 87-88) Grain size: 24.2-26.9 x 17.9-19.3 mm, n = 10. (NY 809; F. W. Pennell 14467; Peru) Monads. Isopolar, prolate (P/E = 1.34), amb Philacra steyermarkii Maguire (Plate 3: 67-69) circular slightly multilobate. Polyzonocolporate (NY 277; J. A. Steyermark 93615) (17 colpori). Ectocolpi long (34.4 mm) and narrow, Monads. Isopolar, prolate (P/E = 1.40), amb endocingulum present (zonorate). Exine 4.0 mm apparently circular-triangular, angulaperturate. Tri- thick, psilate to slightly scabrate, tectate (columellae colporate, zonoaperturate. Ectocolpi long (21.9 mm) visible through the tectum). Nexine thicker than and narrow, constricted in the equatorial region. sexine. Grain size: 45.7-52.7 x 30.8-39.5 mm, n = 6. Ora elliptic, lalongate (3.4 x 7.9 mm), sometimes constricted. Exine 1.6 mm thick, psilate and tectate Rapateaceae (columellae visible through the tectum). Sexine and nexine approximately of the same thickness. Grain Kunhardtia rhodantha Maguire (Plate 1: 1-3) size: 26.2-31.1 x 18.6-22.1 mm, n = 12. (NY 3056; B. Maguire & L. Politi 27954; Cerro Sipapo, Venezuela, Jan. 1, 1949) Poecilandra pumila Steyerm. (Plate 3: 70) Monads. Heteropolar, oblate (P/E = 0.65). Mono- (NY 297; B. Maguire & S. S. Tillett 43800) sulcate, sulcus long (39.8 mm). Exine 1.5 mm thick, Monads. Isopolar, oblate-spheroidal (P/E = 0.99), psilate, tectate. Sexine and nexine approximately of amb circular. Tricolporate, zonoaperturate. Ectocolpi the same thickness. Grain size: 27.4-36.9 x 33.0-56.6 long (21.2 mm) and narrow, ora apparently elliptic mm, n = 10. lalongate, fastigium present. Exine 1.3 mm thick, Comments: Described in Carlquist (1961). psilate and tectate. Sexine and nexine indistinguishable for measuring. Grain size: 27.3-32.2 x 26.6-31.7 mm, n = 10. Rubiaceae

Tyleria spathulata Gleason (Plate 3: 71) Borreria laevis (Lam.) Griseb. (Plate 3: 77-79) (NY 295; B. Maguire et al. 30092) (NY 815; H. H. Smith 87; Colombia) Monads. Isopolar, prolate-spheroidal (P/E = 1.13), Monads. Isopolar, oblate-spheroidal (P/E = 0.94), amb circular slightly lobate. Tricolporate, zonoapertu- amb circular. Polyzonocolpate (9 colpi), short and rate. Ectocolpi long (23.4 mm, n = 3) and narrow (2.0 narrow (5.4 mm long, 0.8 mm wide), ends sharp.

Collectanea Botanica vol. 29 (2010): 31-49, ISSN: 0010-0730, doi: 10.3989/collectbot.2010.v29.004 Tepuian pollen flora (Venezuelan Guayana) 47

Exine 2.7 mm thick, tectate, psilate (columellae turate. Ectocolpi long, sharp ends. Exine 3.8 mm visible through the tectum). Sexine thicker than thick, psilate, tectate. Sexine thicker than nexine. nexine. Grain size: 29.8-36.6 x 30.8-38.5 mm, n = 8. Grain size: 39.7-42.6 x 40.7-40.9 mm, n = 3. Comments: Borreria remota (Lam.) Bacigalupo Comments: Duranta obtusifolia, in Steyermark & E. L. Cabral, in Steyermark et al. (1995-2005). et al. (1995-2005).

Sarraceniaceae ACKNOWLEDGEMENTS

Heliamphora neblinae Maguire var. neblinae The authors are grateful to O. Huber, who noticed the existence (Plate 3: 80-81) of the Maguire pollen collection and encouraged its study. (NY 2736; B. Maguire et al. 37019; Venezuela) The comments of one anonymous reviewer contributed to the Monads. Isopolar, prolate (P/E = 1.41), amb improvement of the manuscript. Funding was provided by the Higher Council for Scientific Research (CSIC) of Spain usually quadrangular, angulaperturate. Tretracolpo- and the Biological Sciences Commission of the Institute for rate, zonoaperturate. Ectocolpi long (20.3 mm) and Catalan Studies (IEC). narrow. Os indistinct, sometimes visible in optical section. Exine 1.3 mm thick, psilate, tectate. Sexine and nexine approximately of the same thickness. REFERENCES Grain size: 30.0-31.2 x 21.4-22.8 mm, n = 3. Comments: Heliamphora minor Gleason de- Berry, P. E. & Riina, R. 2005. Insights into the diversity of the Pantepui flora and the biogeographic complexity of the scribed in Salgado-Labouriau & Villar (1992) and Guayana Shield. Biol. Skr. 55: 145-167. Rull (2003). Briceño, H. O. & Schubert, C. 1990. Geomorphology of the Gran Sabana, Guayana Shield, southeastern Venezuela. Geomorphology 3: 125.141. Ternstroemiaceae Carlquist, S. 1961. Pollen morphology of Rapateaceae. Aliso 5: 39-66. Freziera sericea Humb. & Bonpl. (Plate 3: 63-64) Funk, V. A., Susanna, A., Stuessy, T. F. & Bayer, R. J. (Eds.). (NY 209; 892) 2009. Systematics, evolution, and Biogeography of Com- Monads. Isopolar, prolate (P/E = 1.46), amb circular. positae. IAPT, Vienna. Gorzula, S. & Huber, O. 1992. Consideraciones finales. In: Hu- Tricolporate, zonoaperturate. Ectocolpi long (11.7 mm) ber, O. (Ed.), El Macizo del Chimantá. Un ensayo ecológico and narrow, os indistinct. Exine 0.7 mm thick, psilate. tepuyano. Oscar Todtmann Eds., Caracas: 325-330. Sexine and nexine indistinguishable for measuring. Hokche, O., Berry, P. E. & Huber, O. (Eds.). 2008. Nuevo Grain size: 14.0-15.5 x 9.3-11.2 mm, n = 10. Catálogo de la Flora Vascular de Venezuela. Fundación Instituto Botánico de Venezuela, Caracas. Comments: Eurya sericea (Kunth) Szyszyl., in Howard, R. A. & Boom, B. M. 1990. Bassett Maguire-An Hokche et al. (2008). annotated biography. Mem. New York Bot. Gard. 64: 1-28. Huber, O. 1988. Guayana Highlands versus Guayana Lowlands, a reappraisal. Taxon 37 (3): 595-614. Velloziaceae Huber, O. 1994. Recent advances in the phytogeography of the Guayana region, South America. Mém. Soc. Biogéogr. Barbacenia magalhaesii L. B. Sm. (Plate 1:4-5) 4: 53-63. (NY 438; B. Maguire et al. 49062; Brazil) Huber, O. 1995a. Geographical and physical features. In: Berry, Monads. Heteropolar, oblate (P/E = 0.61). Mono- P. E., Holst, B. K. & Yaskievych, K. (Eds.), Flora of the Venezuelan Guayana 1, Introduction. Missouri Botanical sulcate, sulcus long. Exine < 1 mm thick, perforate. Garden Press, St. Louis: 1-61. Sexine and nexine indistinguishable for measuring. Huber, O. 1995b. Conservation of the Venezuelan Guayana. In: Grain size: 16.9-24.6 x 30.6-36.4 mm, n = 5. Berry, P. E., Holst, B. K. & Yaskievych, K. (Eds.), Flora of the Venezuelan Guayana 1, Introduction. Missouri Botanical Garden Press, St. Louis: 193-218. Verbenaceae Huber, O. 2005. Diversity of vegetation types in the Guayana region: an overview. Biol. Skr. 55: 169-188. Duranta mutisii L. f. (Plate 3: 73-74) Maguire, B., Wurdack, J. J. & Huang, Y. 1974. Pollen grains of some American Olacaceae. Grana 14: 26-38. (NY 1526; W. H. Camp E-5203, Ecuador) Nogué, S., Rull, V. & Vegas-Vilarrúbia, T. 2009. Modeling Monads. Isopolar, spheroidal (P/E = 1.00), amb biodiversity loss by global warming on Pantepui, northern triangular, angulaperturate. Tricolporate, zonoaper- South America. Clim. Change 94: 77-85.

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Punt, W., Hoen, P. P., Blackmore, S., Nilsson, S. & Le Thomas, O. A. (Eds.), Urumaco and Venezuelan palaeontology-the A. 2007. Glossary of pollen and spore terminology. Rev. fossil record of the northern Neotropics. Indiana University Palaeobot. Palynol. 143: 1-81. Retrieved Oct 25, 2010, Press, Bloomington: 84-102. from http://www3.bio.uu.nl/palaeo/glossary Rull, V., Abbott, M. B., Vegas-Vilarrúbia, T., Bezada, M., Rull, V. 2003. An illustrated key for the identification of pollen Montoya, E. Nogué, E. & González, C. 2010. Paleoenvi- from Pantepui and the Gran Sabana (Eastern Venezuelan ronmental trends in Venezuela during the last glacial cycle. Guayana). Palynology 27: 99-133. In: Sánchez-Villagra, M., Carlini, A. A. & Aguilera, O. A. Rull, V. & Vegas-Vilarrúbia, T. 2006. Unexpected biodiversity (Eds.). Urumaco and Venezuelan palaeontology-the fossil loss under global warming in the neotropical Guayana record of the northern Neotropics. Indiana University Press, Highlands. Global Change Biol. 12: 1-9. Bloomington: 52-83. Rull, V. & Vegas-Vilarrúbia, T. 2008. Biopiracy rules hinder Salgado-Labouriau, M. L. & Villar, L. 1992. Contribución conservation efforts. Nature 453: 26. a la flora polínica de los tepuyes. In: Huber, O. (Ed.), El Rull, V., Vegas-Vilarrúbia, T., Nogué, S. & Montoya, E. 2008a. Macizo del Chimantá. Un ensayo ecológico tepuyano. Oscar Bureaucratic obstruction of conservation science in the Todtmann Eds., Caracas: 219-236. Guayana highlands. Conservation Biol. 22: 508-509. Steyermark, J. A., Berry, P. E., Yatshievych, K. & Holst, B. Rull, V., Vegas-Vilarrúbia, T., Nogué, S. & Huber, O. 2008b. K. (Eds). 1995–2005. Flora of the Venezuelan Guayana. Conservation of the unique neotropical vascular flora of Missouri Botanical Garden, St. Louis & Timber Press, the Guayana highlands in the face of global warming. Portland, OR. Conservation Biol. 23: 1323-1327. Tellería, M. C. 2008. Taxonomic significance of pollen Rull, V. 2009. Pantepui. In: Gillespie, R. G. & Clague, D. A. types in the Guayana Highland-centered composite (Eds.), Encyclopedia of Islands. University of California genera of Mutisioideae (Asteraceae). Bot. J. Linn. Soc. Press, Berkeley: 717-720. 156: 327-340. Rull, V. 2010. The Guayana Highlands: a natural laboratory for Ubiergo, P., Lapp, M. & Torrecilla, P. 2009. Morfología del polen the biogeographical and evolutionary study of the neotropi- de especies de Gongylolepis (Mutisieae: Asteraceae) de la cal flora. In: Sánchez-Villagra, M., Carlini, A. A. & Aguilera, Guayana venezolana. Anales Jard. Bot. Madrid 66: 93-107.

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Appendix. List of species/slides of the Maguire collection analyzed, which pollen grains are unsuitable for a morphological study.

Acanthaceae Justicia carthaginensis Jacq. (NY 846; O. L. Haught 4753; Colombia)

Annonaceae Duguetia riberensis Aristeg. ex Mass & Boon (NY 658; L. Aristeguieta 5308; S. America)

Boraginaceae Cordia leucophylctis Hook. f. (NY 862; Chapin 1125; Indefatigable Is., Galapagos)

Cyatheaceae Cyathea arborea (L.) Sm. (NY 2995; E. L. Little, Jr., 22 June 1950; El Verde, Puerto Rico)

Dennstaedtiaceae Hypolepis repens (L.) Presl. (NY 1546; Cuatrecasas 16092; Colombia)

Dryopteridaceae Diplazium lindbergii (Mett.) Christ (NY 1524; Holdridge 1608; Ecuador)

Fabaceae Inga aff. punctata Willd. (NY 769; G. Tessmann 3443; Peru)

Lamiaceae Hyptis capitata Jacq. (NY 754; Toro 44; Colombia)

Myrsinaceae Cybianthus brownii Gleason (NYBG 2363; Maguire 32104; Brit Guiana)

Myrtaceae Eugenia florida D. C. (NY 2170; H. S. Irwin et al. 55206; Suriname)

Ochnaceae Elvasia quinqueloba Spruce (# 34812; B. Maguire & J. J. Wurdack, 287)

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