Lianas of the Guianas

A guide to woody climbers in the tropical forests of Guyana, Suriname and French Guiana

Bruce Hoffman, Ph.D. and Sofie Ruysschaert

Lianas of the Guianas: A guide to woody climbers in the tropical forests of Guyana, Suriname and French Guiana Bruce Hoffman, Ph.D. Sofie Ruysschaert

LM Publishers Netherlands

Contributions by Mark J. Plotkin, Ph.D. Frits van Troon Joeri Zwerts

Disclaimer The information provided within this book is for general informational purposes only. While we try to keep the information up-to-date and correct, there are no representations or warranties, express or implied, about the completeness, accuracy, reliability, suitability or availability with respect to the information, products, services, or related graphics contained in this book for any purpose.

Cover and graphic design / NANCY VALIES

Icon art Kristian Johnson Michiels

Image editing and print High Trade BV, Zwolle

ISBN: 98709780978908

A Word from the Sponsor In a country like Suriname with a pristine tropical rainforest, it is important to be aware of the richness of the forest. Lianas are very well known, but it was surprising to read about the number and types of liana species in our country. It is therefore with much pleasure that we have agreed to sponsor the printing of this publication. It is our hope that this reference work shall be well received by the general public and especially our youth. We hope they will gain much knowledge on the subject of lianas. We thank and congratulate everyone who has given their utmost to put this reference work together.

The Suriname Conservation Foundation Henna J. Uiterloo LLB Executive Director

With the generous support of Amazon Conservation Team Alberta Mennega Stichting (Netherlands) Stichting het van Eeden Fonds (Netherlands) Hugo de Vries Fonds (Netherlands) Naturalis Biodiversity Center (Netherlands) LM Publishers

A guide to woody climbers in the tropical forests of Guyana, Suriname and French Guiana

Bruce Hoffman, Ph.D. and Sofie Ruysschaert with contributions by Mark J. Plotkin, Ph.D., Frits van Troon, and Joeri Zwerts

FOREWORD

Schlegelia violacea (Schlegeliaceae), with flowering branch extending down from host tree.

FOREWORD According to the eminent Neotropical botanist Thomas Croat, the presence of lianas is the single most important physiognomic feature that distinguishes tropical from temperate forests. This distinction holds true in the collective mind of the general public as well: ever since Johnny Weissmuller’s Tarzan first swung through the jungle, we all associate woody vines with tropical forests.

Because they often flower and fruit in the canopy far above the forest floor, lianas are notoriously difficult to collect and study. Famed 20th-century ethnobotanist Richard Schultes searched seven years before he was able to find the fabled ayahuasca vine in flower, and he then had to fell seven trees to access the blossoms, so interwoven was the liana in the crowns of these forest giants. Challenges like these long hindered the study of these tropical vines by non-indigenous scholars, hence the likely immediate worth of any published research that attempts to digest their enormous diversity for both academic and lay readers.

Recent years have seen a plethora of field guides devoted to explaining and identifying tropical organisms: palms, birds, reptiles, freshwater fishes and others. With such manuals in hand, both the amateur and the professional biologist can walk through these forests and identify flora and fauna as never before. However, prior to the herculean effort leading to this publication, a regional guide to these challenging and mysterious life forms has not been available.

The race is on to better understand tropical rainforests and their component species in the face of relentless deforestation and extractive pressures. It is my great hope that this pioneering research by my longtime associate Dr. Bruce Hoffman and his local colleagues—fellow scientists and indigenous mentors—will make a significant contribution to our better valuing, utilizing and protecting these magnificent species and the rainforests in which they thrive.

FOREWORD

Dr. Mark J. Plotkin

Lianas of the Guianas

7

LIA N A S OF T H E G UI A N A S Guide

1

BRUCE HOFFMAN AND JOERI ZWERTS

INTRODUC TION Overview of the Guide

The front of the guide consists of three brief chapters: a general introduction to climbing plant diversity and the Guianas, an illustrated botanical glossary, and a `how-to-use’ chapter with tables of the icons used in family chapters and other identification aids. A vegetative key to liana families and genera is in development but could not be included in this volume. The core of the book includes forty-four family chapters in alphabetical order, with 175 genera and 384 species in total. Detailed descriptions are provided for all documented families and genera with woody climbers and approximately 38% of the 1,023 species. Scientific names are based upon the modern `APG III’ classification (Bremer et al.

2009) and the Taxonomic Names Resolution Service v. 4.0. (TNRS 2016). In the appendices, readers will find a comprehensive checklist of herbaceous and woody climbers in the Guianas (excluding ferns), text-based botanical glossary, common names index, bibliography, and author image credits.

We hope that this guide will serve as a useful tool for pragmatic purposes such as forest inventories, sustainable logging, ecotourism, non-timber forest product development and scientific research. We also hope that it will be an enjoyable book for those simply wanting to discover more about these fascinating, iconic life forms of tropical forests.

Lianas of the Guianas

INTRODUCTION - OVERVIEW OF THE GUIDE

Lianas (woody vines) and other high-climbing plants, experienced in tropical forests mostly as mysterious bare woody stems disappearing into the canopy, are notoriously difficult to observe, document, and identify. The aim of this guide is to provide readers with a greater understanding and (hopefully) appreciation of liana diversity, ecology, and related human significance in the Guianas, and thereby increase knowledge and appreciation of the region’s tropical forests as a whole. The intended audience is broad, including students, scientists, resource managers, forest inventory teams, international and national institutions, tourists, forest-dependent communities and the general public. The information in this guide was compiled from a wide variety of sources, including empirical research of the authors and colleagues, herbarium specimens, flora and field guides, scientific websites and databases, forestry specialists, ethnobotanical references, indigenous and tribal peoples, illustrators and photographers.

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The Significance of Lianas Climbers are species that require the support of another plant or surface for growth during at least one part of their life cycle. The climbing habit occurs throughout the world with an increasing abundance towards the equator and within lowland tropical forests (Gentry 1991). Woody vines, commonly known as lianas, are iconic symbols of tropical forest ecosystems. Using active climbing mechanisms such as twining shoots, tendrils or adhesive roots and passive mechanisms such as spines, rough surfaces, and angled branches, lianas take advantage of the biologically-expensive, vertical structure of trees to gain relatively inexpensive access to the light-rich canopy. The resources that are thus conserved can be invested elsewhere to gain a competitive advantage in growth and reproduction. Once in the canopy, lianas often extend horizontally and may produce more leaf area than the crowns of the trees they occupy (Putz 1983).

The success of the climbing strategy is revealed by the widespread occurrence of climbing species in unrelated plant groups. Aside from a rich representation among angiosperms (flowering plants), climbers can also be found amongst neotropical palms (Desmoncus), ferns, orchids, and gymnosperms (Gnetum). In the Neotropics, three distantly-related angiosperm plant families typically dominate in species diversity, including the Bignoniaceae, Sapindaceae and Leguminosae (Fabaceae). The evolution of the climbing habit in forests may have resulted from the diverse surfaces and spaces presented by multi-story forest canopies, interactions with diverse pollinators and seed dispersers, or the obvious adaptive benefit of escaping the shaded forest floor. On average, woody climbers compose around 25% of the woody stems and species in rainforests, forming a key element in forest diversity, structure and composition (Schnitzer, et al. 2012, Gentry 1991).

INTRODUCTION - OVERVIEW OF THE GUIDE

Lianas play an important role in forest ecology, gap dynamics and forestry operations. Dormant seeds, suppressed seedlings and vegetative sprouts from fallen lianas, in combination with high growth rates, result in the rapid proliferation of lianas in forest gaps. For trees, lianas can impede growth rates and regeneration, and increase mortality. To minimize damage to commercial species, lianas are often cut from trees by foresters before felling. Due to their narrow stems and low wood density, lianas sequester relatively little carbon compared to the trees whose growth and turnover rate they influence negatively.

Therefore the net carbon sequestration of a forest becomes lower with increased liana occupation (Laurance 1997). Lianas appear to be out-competing trees and becoming more abundant in the face of global warming and increasing seasonality in some tropical forests (Phillips, et al. 2002).

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Throughout the tropical world, forests and their lianas provide a bounty of ecosystem and economic services in the form of food, raw materials, water, oxygen, medicine, biodiversity, tourism and carbon sequestration, to name but a few. In the canopy, lianas grow laterally and establish important canopy pathways for animals to move through the forest. The negative impact of lianas upon tree growth is counterbalanced by the horizontal lattice that helps keep trees from falling over during storms. Lianas are also an important part of the food web providing leaves, flowers, fruits, nectar, water as well as nesting material and habitat. To maintain these services, it is important to understand the system from which they are derived, a system in which climbing plants play an unmistakable role in a complex ecological web of direct and indirect interactions.

Lianas of the Guianas

The Guianas Region This guide focuses upon woody climbers native to three territories in northeastern South America: French Guiana, Guyana, and Suriname. Guyana (formerly British Guiana) and Suriname (formerly Dutch Guiana) are independent countries, while French Guiana remains an overseas department of France. These three culturally diverse and unique territories are known collectively as “the Guianas”, although the underlying Precambrian rock of the Guiana Shield formation extends over a much greater area, including Brazil, Venezuela and eastern Colombia. The greater region is associated with ancient, nutrient-poor soils, translucent ‘black water’ creeks, savannas and relatively low diversity forests compared to western Amazonia. The western Guiana Shield, including parts of Guyana, has sandstone tepui plateaus to 3000 meters, spectacular waterfalls, and a highly endemic flora (Steyermark et al. 1995). The eastern Guiana Shield (French Guiana, Suriname) is less precipitous, with broad expanses of lowland tropical forest interrupted by scattered mountain ranges, granitic domes and a single tepui reaching 1300 meters. From European colonial times to the present, population growth and development in the Guianas region has remained concentrated in accessible areas along the Atlantic coast and major rivers.

Although unsustainable extractive activities such as gold-mining are encroaching, the interior forests of the Guianas have been largely spared by difficulty of access, and the region composes one of the world’s largest remaining contiguous expanses of tropical forest. Human presence in the inland forests consists largely of proud indigenous and tribal (Maroon) forest communities, with largely subsistence-based livelihoods.

INTRODUCTION - OVERVIEW OF THE GUIDE

As revealed in the species accounts of this guide, lianas and other climbers are ubiquitous across the Guianas in a wide variety of habitats. Sun-loving, slender lianas thrive in open areas and forest edges, while shade-tolerant, robust lianas are abundant in old growth forests. This guide is likely to be useful over a much greater area than the Guianas, as many of the described taxa occur in adjoining areas and in the Neotropics at large.

Map of the Guianas region, including Guyana, Suriname, and French Guiana

Lianas of the Guianas

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Climbers of the Guianas: Diversity, Growth Forms, Unique Characters

INTRODUCTION - OVERVIEW OF THE GUIDE

Climbing plants in the tropics and the Guianas exhibit great variety, including high taxonomic diversity (many families and species), diverse growth forms, and diverse morphological adaptations for a climbing or crawling way of life. Even species of the same family or genus may exhibit very different climbing solutions, and some species opportunistically switch from self-supporting to a climbing form when the opportunity arises. We estimate 59 plant families and 253 genera of the Guianas to include at least one climbing species. Figure 2 shows the species richness and distribution of climber growth forms (both woody and herbaceous) within 21 plant families. Some families represent only one climbing growth form (Bignoniaceae, Clusiaceae), while others have multiple climbing growth forms among their species (Apocynaceae, Convolvulaceae). In this comparison, the non-climbing species were excluded (e.g., Leguminosae trees). The Apocynaceae, Bignoniaceae and Leguminosae families have the greatest number of climbing species, with 123, 103 and 96 species respectively. However, the dissimilar, herbaceous monocot family Araceae has the fourth largest number of climbing species at 77. In the following few pages we provide an overview of common climber growth forms.

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Figure 2. The number of species and growth form distributions among the 21 most climber-rich plant families in the Guianas Climber Checklist. Growth form definitions: CS, climbing shrub; HE-L, liana-like hemi-epiphyte; HE-T, tree-like hemi-epiphyte; HE-V, vine-like (herbaceous) hemi-epiphyte; L, liana woody, slender to robust; L-SW, sub-woody lianas; V, vines (herbaceous).

Lianas of the Guianas

Lianas (L) are climbers that are rooted in the soil, have a main stem, require support at some point in their life cycle and produce at least some tissues with wood. Stem diameters range from less than 0.5 cm diameter to greater than 30 cm diameter. Lianas of stem diameter > 0.5 cm are the main focus of this book as they are the most widely-dispersed, locally abundant and ecologically significant forest climbers. Climbers with fine “woody” stems less than 0.5 cm diameter can be considered perennial vines or vine-like lianas (L-V).

2

3

4

INTRODUCTION - OVERVIEW OF THE GUIDE

The stem tissues of many forest climbers exists somewhere along a continuum between ‘woody’ and ‘herbaceous’. Sub-woody lianas (L-SW), including many Cucurbitaceae and Vitaceae species, often have robust, green stems that appear to produce a soft wood. Desmoncus is a liana genus in the palm family that, like all monocots, does not have “true wood”. Yet Desmoncus stems are tough, and they are included as lianas in this guide.

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1. Pyrostegia venusta (Bignoniaceae), with orange tubular flowers, lobes recurved at maturity [BH] 2. A robust liana, Croton pullei [BH] 3. Petrea bracteata [BH] 4. Gurania reticulata [PT] 5. Subwoody liana stem with tendril, Cayaponia sp. (Cucurbitaceae) [PT]

Lianas of the Guianas

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INTRODUCTION - OVERVIEW OF THE GUIDE

Climbing shrubs (CS) are woody, terrestrial plants that lack a single main stem and use spreading branches to lean and climb, often rather clumsily, over other plants. They can be aided in their attachment by hooks and spines. Some Gouania (Rhamnaceae), Piper (Piperaceae) and Randia (Rubiaceae) species fit this description. Some canopy-occupying shrubs and cactus species occupy a growth form gray area somewhere between epiphytes, hemi-epiphytes and lianas.

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6

7

8

9

Lianas of the Guianas

Vines (V) are similar in many respects to lianas, but lack wood and generally remain lower in the canopy or in open areas. The flowers of vines are often large and showy. Two vine-rich plant families in the Guianas are the Apocynaceae and the Convolvulaceae. Conspicuously non-woody, slender vines are not included in this guide. Definitions can be tricky because the woody stems of lianas and basally woody climbers commonly produce herbaceous shoots (e.g., Mendoncia species).

10

11

INTRODUCTION - OVERVIEW OF THE GUIDE

12

6. Lomariopsis japurensis (Lomariopsidaceae), a common climbing fern in the Guianas [BH] 7. Hylocereus scandens, (Cactaceae) epiphytic or climbing cactus [PT] 8. Piper hispidum, (Piperaceae) herb or shrub, sometimes weakly climbing [PT] 9. Randia armata (Rubiaceae), climbing shrub with spines [PT] 10. Dichorisandra hexandra (Commelinaceae), a weak, herbaceous climber [BH] 11. Ipomoeae batatoides (Convolvulaceae), a vine or very slender liana [PT] 12. Dioscorea amazonum (Dioscoreaceae), perennial vine with large tubers [BH]

Lianas of the Guianas

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‘Hemi-epiphyte’ is a term for plants that live perched upon trees like (epiphytic) orchids, yet also have an aerial root connection to the ground, unlike most orchids. Three types of hemiepiphytes are recognized in this guide.

Tree-like “primary” hemi-ephiphytes (HE-T), including Clusia (Clusiaceae), Coussapoa (Urticaceae), Ficus (Moraceae) and Oreopanax (Araliaceae) species, establish as robustly woody epiphytes in the canopy and then extend roots to the forest floor. Strangler figs, comprising many species, are well known for engulfing and killing host trees in the tropics.

Vine-like “secondary” hemi-epiphytes (HE-V), including herbaceous Araceae and Cyclanthaceae species, and the Vanilla orchid genus, begin life on the forest floor and climb up tree trunks with clasping roots to reach higher light conditions. Heteropsis, commonly known as cipó, kamina, or nibbi, is a common green-stemmed hemi-epiphyte on tree trunks that produces extremely strong, pencil-thick, woody roots.

INTRODUCTION - OVERVIEW OF THE GUIDE

Liana-like hemi-epiphytes (HE-L) can be similar to either HE-V or HE-T species. They are robustly woody to sub-woody and may begin life on the ground or in the canopy. Representatives include species of Adelobotrys (Melastomataceae), Schlegelia (Schlegeliaceae), Schradera (Rubiaceae), Ericaceae (Cavendishia, Satyria), Marcgraviaceae (Marcgravia, Marcgraviastrum, Souroubea) and Solanaceae (Markea, Solandra). All HE types are included in the Guianas climber checklist, but only HE-L taxa are covered in the family chapters.

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13

14

15

16

Lianas of the Guianas

True epiphytes are plants that never root in the soil. They complete an entire life cycle anchored to a host plant, upon which they germinate as seeds or propagules and become rooted. The majority of epiphytes in the Guianas belong to the Orchidaceae, Bromeliaceae and fern families. Parasitic plants extract water and nutrients from a ‘host’ plant upon which they completely depend for their survival.

Many species in the Loranthaceae family (fowru doti) are parasites that occupy tree crowns.

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18

13. Clusia grandiflora (Clusiaceae), a tree-like hemi-epiphyte with woody aerial roots [BH] 14. Clusia palmicida, rooted at host tree base [BH] 15. Heteropsis flexuosa (Araceae), a vine-like hemi-epiphyte with woody aerial roots (upper left in photo) [BH] 16. Philodendron solimoesense (Araceae), a vine-like hemi-epiphyte with aerial roots [BH] 17. Vanilla planifolia (Orchidaceae), a vine-like hemi-epiphyte in the orchid family [BH] 18. Guzmania lingulata (Bromeliaceae), a true epiphyte - perches upon host trees or other surfaces without extending aerial roots to the soil [PT]

Growth form (habit or habitus)

Code

No. species (1,307 total)

Non-woody

Vines and creepers

V

90 (7%)

Sub-woody

Lianas, soft or woody only at base

L-SW

55 (4%)

Vine-like hemi-epiphytes

HE-V

103 (8%)

Lianas, slender but woody, < 0.5 cm diam.

L-V

174 (13%)

Lianas, small to robust, > 0.5 cm diam.

L

727 (56%)

Liana-like hemi-epiphytes

HE-L

44 (3%)

Tree-like hemi-epiphytes

HE-T

88 (7%)

Clambering or climbing shrub

CS

26 (2%)

Woody

In Table 1, we provide species counts for six climber growth forms in the Guianas climber checklist, regardless of family. Small to

INTRODUCTION - OVERVIEW OF THE GUIDE

Table 1. Number of species by growth form in the Guianas climber checklist (Appendix 2).

robust lianas are the dominant growth form (56%).

Lianas of the Guianas

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T H E E T H N O B OTA N Y of Lianas

Korotai, a Trio shaman in Kwamalasamutu village, Suriname, drinking medicinal water from the cut stem of a liana - Doliocarpus dentatus (Dilleniaceae), also known as kapadula (GU), sakĂŤtaitu (Tr), and watra titĂŠ (Sr).

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1

THE ETHNOBOTANY OF LIANAS Western botanists have often deemed lianas too difficult to access, leaving them in the forest rather than collecting them for the herbaria. Indigenous peoples in these same forests, however, can often distinguish lianas by the appearance of the stem or the fragrance of the bark. Their sophistication at identifying these species is matched only by their astonishing ability to make full use of them. In tropical South America, indigenous peoples have employed the components of lianas as food, arrow and fish poisons, cordage, dyes, stimulants, hallucinogens, euphoriants, and sources of potable water.

Documented western encounters with indigenous botanical knowledge of the Guianas trace back at least to the early 18th century, notably in the writings of Edward Bancroft, a sailor, ethnobotanist, physician, business tycoon, American diplomat and double agent for the British Crown. Born poor in western Massachusetts in 1735, Bancroft went to sea as a teenager in search of fame, fortune and adventure. History then records him practicing medicine (seemingly without having attended medical school) in what was then British Guiana. Based on his experience and observations, in 1769 he published a book entitled “Essay on the Natural History of Guiana” that contained numerous accounts of local plants, animals and peoples, including a thirty-page discussion of curare. Bancroft noted that each Amerindian tribe had its own recipe for arrow poison, and he recorded the curare of the Akawaio tribe of northwestern Guiana—perhaps the first detailed account of the deadly poison three decades before Alexander von Humboldt famously observed the process in neighboring Venezuela.

Ethnobotanical studies will undoubtedly reveal further intriguing applications for rainforest lianas, if they can be conducted before the tide of western culture sweeping through Amazonia obliterates such indigenous knowledge. By way of example, approximately thirty percent of the Trio indigenous ethnopharmacopeia in Suriname is derived from liana species. To the few living scientists who have directly immersed themselves in learning at the feet of indigenous botanical scholars of the Guianas— Dr. Bruce Hoffman and I both count ourselves among their scant ranks—the value of perpetuating the regional shamans’ knowledge and their habitats could not be more unambiguous.

Dr. Mark J. Plotkin

Lianas of the Guianas

THE ETHNOBOTANY OF LIANAS

More prominently, in 1804, the naturalist Charles Waterton set sail for British Guiana to oversee his family’s plantations near Georgetown. After eight years near the coast, he departed for the interior, in search of what was said to be the deadliest curare of the Guianas, that made by the Macushi tribe.

Waterton’s expedition—which he conducted barefoot, and in the rainy season—was successful. Unlike the poison collected by Bancroft, which was made from Strychnos guianensis, the seemingly more toxic Macushi curare was prepared from the liana now known as Strychnos toxifera, and confirmed by the far more meticulous Schomburgk brothers who traveled in Waterton’s footsteps several decades later. Waterton brought samples back to England, and facilitated a number of experiments that eventually led to the use of Amazonian curares as anesthetics in abdominal surgery.

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I L LU S T R AT E D G LO S S A R Y

Unique liana structures This chapter provides labeled illustrations for structures unique to climbers and for flowering plants in general.

Climbing mechanisms All climbers can be categorized by one or more climbing mechanisms. This guide groups mechanisms within five major categories (See following illustrations and icon tables for climbing mechanisms in the next chapter): a) Tendril climbers – Tendrils are long, slender organs developed from modified leaves, stems or branches to clasp surrounding structures, providing an elastic and robust form of attachment. The location, shape (3D spiral, 2D coil) and number of tendrils are important in plant identification. b) Twining climbers - Twining plants revolve their shoots in a spiral motion as a means of exploration, attaching to objects and climbing. Some plants have tendril-like shoots or leaves that grasp objects instead of spiraling around them (e.g. Salacia, Securidaca). c) Hook/Spine/Hair climbers – Plants use this mechanism to hold onto the host tree with downward pointing adhesive hairs, rough surfaces, hooks, or spines. The spines are often paired and represent modified stipules. d) Clambering branch and petiole climbers – This method involves sharply angled branches or elbow-like or twisted petioles. Both hook and branch climbing are passive methods, and plants are more likely to lose their grip compared to twiners, tendril climbers, and root climbers. e) Root climbers – These plants cling to a support with aerial roots that emerge from the stem. They grow into irregular surfaces or attach themselves with sticky glandular secretions. This mechanism requires close contact with the host tree’s main trunk, and climbers typically occupy only lower levels of the canopy. Most root climbers are vine-like or liana-like secondary hemi-epiphytes.

Stem anatomy

ILLUSTRATED GLOSSARY

See woody cross-section pattern icons here and icon tables in the next chapter.

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Flexibility in lianas is due partly to their slender stems, but also results from their shapes and internal anatomy. Liana stems often have fewer fibers and an abundance of large-diameter, water-conducting xylem vessels (wood). This allows them to pipe more water and thus produce more leaf area than might be expected for their stem size. The xylem and phloem (sugar transport) tissues of liana stems are often uniquely arranged (compared to trees), a condition sometimes termed ‘anomalous growth’ or a ‘cambial variant’ (Carlquist 1991, Pace et al. 2009). With the vascular tissues spread out within the stem, tree fall damage is less likely to be fatal to the liana, because even if a stem is split in half, there should still be some functioning cables to deliver sugars, nutrients and water. This unique anatomy often results in unique patterns visible to the naked eye which can be useful in identification. However, we request that users of this guide not destroy lianas just to observe the stem cross-section. A lengthwise slash or cutting of a small side branch is usually sufficient to reveal the internal anatomical pattern.

Lianas of the Guianas

Woody Stem cross-section patterns icons

Eccentric Simple

Foraminate/specks (discrete)

Foraminate/islands (discrete)

Concentric rings (bands narrow or broad) Sometimes flattened but still centered

Lobed-1

Lobed-2 Divided (poly-stelic)

Radial

ILLUSTRATED GLOSSARY

Compound (poly-stelic)

Interrupted (Bignoniaceae)

Lianas of the Guianas

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1. C LIMBING MECHANISM - HOOK CLIMBERS

Rubiaceae - Uncaria Paired tendril-hooks or spines from modified axillary stipules (coiled 2D or straight)

Loganiaceae - Strychnos

Arecaceae - Desmoncus

Woody tendril-hook from modified axillary

Barbed, reflexed hooks from modified leaflets at tail-like apex

stipules paired or solitary (coiled 2D)

of palm leaf

1. C LIMBING MECHANISM - TENDRILS

Passifloraceae - Passiflora

Curcurbitaceae

Vitaceae - Cissus

Tendril at petiole base (spiral)

Tendril at 90 degrees to

Tendril opposite petiole base (spiral)

ILLUSTRATED GLOSSARY

petiole base (spiral)

Bignoniaceae

Sapindaceae

Smilacaceae - Smilax

Tendril a modified leaflet on a compound leaf

Long, forked tendril from a modified

Paired tendril from opposite

(simple or 2-3 branched, rarely many branched)

flowering branchlet (coiled 2D or spiral)

sides of petiolar sheath (spiral)

Illustrations redrawn with permission from Bobbi Angell illustrations

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Lianas of the Guianas

1

1. C LIMBING MECHANISM - TENDRILS

Leguminosae - Bauhinia Woody tendril-hook from a modified branchlet (coiled 2D)

ILLUSTRATED GLOSSARY

2. L EAVES - SCHEMATIC

Lianas of the Guianas

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ILLUSTRATED GLOSSARY

2. L EAVES - SHAPE

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Awl shaped

Cordate

Cuneate

Deltoid

Elliptic

Lanceolate

Linear

Oblanceolate

Oblong

Obovate

Orbiculate

Oval

Ovate

Palmate Lobed

Pinnate Lobed

Spatulate

Lianas of the Guianas

2. L EAVES - APEX

Acute

Attenuate

Cuspidate

Emarginate

Long tapering

Mucronate

Notched

Obtuse

Rounded

Spine tipped

Truncate

ILLUSTRATED GLOSSARY

Acuminate

Lianas of the Guianas

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